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PALAEONTOLOGIA AFRICANA PALAEONTOLOGIA AFRICANA Volume 47 Annals of the Bernard Price Institute for Palaeontological Research December 2012 ISSN 0078-8554 Supported by PALAEONTOLOGICAL SCIENTIFIC TRUST

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Page 1: PALAEONTOLOGIA AFRICANA

PALAEONTOLOGIA

AFRICANA

PALAEONTOLOGIA

AFRICANA

Volume 47

Annals of theBernard Price Institute

forPalaeontological Research

December 2012

ISSN 0078-8554

Supported byPALAEONTOLOGICAL SCIENTIFIC TRUST

Page 2: PALAEONTOLOGIA AFRICANA

SCHOOL OF GEOSCIENCESBERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCH

Academic StaffDirector and Chair of PalaeontologyB.S. Rubidge BSc (Hons), MSc (Stell), PhD (UPE)Deputy DirectorM.K. Bamford BSc (Hons), MSc, PhD (Witwatersrand)Senior Research OfficersF. Abdala BSc, PhD (UNT, Argentina)L.R. Backwell BA (Hons), MSc, PhD (Witwatersrand)J. Choiniere BSc (Massachusetts), PhD (GWU).Collections ManagerB. Zipfel NHD Pod., NHD PS Ed. (TWR), BSc (Hons)(Brighton), PhD (Witwatersrand)

Post Doctoral FellowsD. Marchi MSc, PhD (Pisa, Italy)V. Fernandez MSc (Lyon 1, France), PhD (Paris VI, France)C.A. Phillips BSc (Hons) (Bournemouth), MSc (OxfordBrookes), PhD (Cambridge)

Editorial PanelM.K. Bamford: EditorL.R. Backwell: Associate EditorB.S. Rubidge: Associate EditorConsulting EditorsDr J.A. Clack (Museum of Zoology, University ofCambridge, Cambridge, U.K.)

Dr H.C. Klinger (South African Museum, Cape Town)Dr K. Padian (University of California, Berkeley,California, U.S.A.)

Dr K.M. Pigg (Arizona State University, Arizona, U.S.A.)Prof. L. Scott (University of the Free State, Bloemfontein)Dr R.M.H. Smith (South African Museum, Cape Town)

Technical and Support StaffPrincipal TechnicianW.P. GermishuizenSenior Administrative SecretaryS. Sejake

Assistant Research TechnicianC.B. Dube

Technicians/Fossil PreparatorsP. ChakaneS. JirahP.R. MukanelaG. NdlovuT. NemavhundiS. Tshabalala

Honorary Staff

Honorary Research AssociatesK. Angielczyk BSc (Univ. Michigan, Ann Arbor), PhD(University of California, Berkeley)

P.J. Hancox BSc(Hons), MSc, PhD (Witwatersrand)R. Reisz BSc, MSc, PhD (McGill University, Montreal)C.A. Sidor BSc (Trinity College) MSc, PhD (University ofChicago)

INSTITUTE FOR HUMAN EVOLUTION

Academic StaffDirectorJ.F. Thackeray BSc (Hons), MSc (UCT), MPhil, PhD (Yale)ReaderL.R. Berger BA (Hons) (GA Southern), PhD (Witwaters-rand)

Research OfficersK. Carlson BSc (Michigan, Ann Arbor), MSc, PhD(Indiana, Bloomington)

J. Kibii BSc, (Nairobi), MSc, PhD (Witwatersrand)B. Kuhn BSc (Washington State), MSc (UCL), PhD (UP)Research AssociatesC. Henshilwood BA (Hons) (UCT), PhD (Cambridge)L. Wadley MSc (UCT), PhD (Wits)Administrative and Support StaffSenior Administrative SecretaryEvlyn HoPreparatorsT. DingiswayoT. MakheleN. Molefe

A. MollepolleS. MotsumiL. SekoweM. Seshoene

Honorary Staff

Honorary Research AssociatesDr Shaw Badenhorst, Transvaal MuseumProf. Steve Churchill, Duke University, USAProf. Francesco d'Errico, BordeauxUniversity, FranceProf. Daryl de Ruiter, Texas A&M University, USAProf. Katerina Harvati, University of Tubingen, GermanyProf. Jacopo Moggi-Cecchi, Laboratori di Antropologia,Dipartimento di Biologia Animale e Genetica, Universitàdi Firenze, Italy

Prof. Peter Schmidt, Zurich, SwitzerlandProf. Himla Soodyall, Human Genomic Diversity andDisease Research Unit (HGDDRU), South AfricanMedical Research Council in conjunction with theNational Health Laboratory Service and University ofthe Witwatersrand

Dr Peter Ungar, University of Arkansas, USA

Page 3: PALAEONTOLOGIA AFRICANA

Articles

Kennedy, W.J. & Klinger, H.C. — Cretaceous faunas from Zululand and Natal,South Africa. The ammonite genus Codazziceras Etayo-Serna, 1979

Kennedy, W.J. & Klinger, H.C. — The ammonite genus Diaziceras Spath, 1921,from the Campanian of KwaZulu-Natal, South Africa, and Madagascar

Kennedy, W.J. & Klinger, H.C. — Cretaceous faunas from Zululand and Natal,South Africa. A new species of the ammonite genus Salaziceras Breistroffer,1936, from the Lower Cenomanian Mzinene Formation

Geraads, D., El Boughabi, S. & Zouhri, S. — A new caprin bovid (Mammalia)from the late Miocene of Morocco

Odes, E.J. & Thackeray, J.F. — Morphometric analysis of modern humancrania: a framework for assessing early Pleistocene hominids

Abstracts16th Biennial Conference of the Palaeontological Society of Southern Africa

Standard Bank/PAST First Phillip Tobias Memorial Lecture

Preface to Prof. Nina Jablonski’s public lecture on “Skin: Its Biology in Black and White”by A. Leenen and R.J. Blumenschine

Jablonski, N.G. — Skin: Its Biology in Black and White

1

3

15

19

25

29

61

62

Volume 47, December 2012

PALAEONTOLOGIA AFRICANA

CONTENTS

ANNALS OF THE BERNARD PRICE INSTITUTE FOR PALAEONTOLOGICAL RESEARCHUNIVERSITY OF THE WITWATERSRAND

ISSN 0078-8554

Page 4: PALAEONTOLOGIA AFRICANA

© 2012BERNARD PRICE INSTITUTE

forPALAEONTOLOGICAL RESEARCH

School of GeosciencesUniversity of the Witwatersrand

Johannesburg

ACKNOWLEDGEMENTS

The Bernard Price Institute for Palaeontological Research gratefully acknowledges financial support for itsprogrammes by

THE COUNCIL’S RESEARCH COMMITTEE, UNIVERSITY OF THE WITWATERSRAND

NATIONAL RESEARCH FOUNDATION (NRF)

DEPARTMENT OF SCIENCE AND TECHNOLOGY (DST)

and the

PALAEONTOLOGICAL SCIENTIFIC TRUST (PAST)

for publication of this journal

Pre-press production by Isteg Scientific Publications, Irene

Printed in South Africa by Ultra Litho (Pty) Ltd, Heriotdale, Johannesburg

Page 5: PALAEONTOLOGIA AFRICANA

Cretaceous faunas from Zululand and Natal,South Africa‡. The ammonite genus

Codazziceras Etayo-Serna, 1979William James Kennedy1* & Herbert Christian Klinger2*

1Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PW, andDepartment of Earth Sciences, South Parks Road, Oxford OX1 3AN, United Kingdom

2Natural History Collections Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa

Received 6 August 2012. Accepted 22 October 2012

INTRODUCTIONThe collections of the Iziko South African Museum

include a small ammonite, just over 50 mm in diameter,from the western shores of False Bay, Lake St Lucia, innorthern KwaZulu-Natal. It is described below as a newspecies of the genus Codazziceras Etayo-Serna, 1979,known previously with certainty only from the typespecies Codazziceras scheibei (Riedel, 1938) (which is ajunior synonym of Ammonites ospinae Karsten, 1858),and C. fina Etayo-Serna, 1979, from the Coniacian ofColombia. Yubariceras gosavicum Wiedmann, 1979, fromthe Coniacian of Austria may also belong to the genus.The present record is thus the first from the SouthernHemisphere.

SYSTEMATIC PALAEONTOLOGY

Suborder Ammonitina Hyatt, 1889Superfamily Acanthoceratoidea de Grossouvre, 1894Family Acanthoceratidae de Grossouvre, 1894Subfamily Euomphaloceratinae Cooper, 1978

Genus Codazziceras Etayo-Serna, 1979

Type speciesLyelliceras scheibei Riedel, 1938, p. 55, pl. 9, figs 7, 8; pl. 13,

fig. 17, by the original designation of Etayo-Serna (1979,p. 83) = Ammonites ospinae Karsten, 1858, p. 110, pl. 4,fig. 3.

Diagnosis‘Small, very evolute, serpenticone, whorl section square

to rectangular on phragmocone, rounded on body chamber.Innermost whorls smooth and constricted, constrictionspersist into early part of ribbed stage. Phragmocone andearly body chamber with strong ribs branching from

umbilical bullae or not or intercalated. Primary ribs withweak inner umbilical tubercles that may be indistinct atsome stage and strong outer umbilicals; all ribs with innerand outer ventrolateral and siphonal tubercles. All tuber-cles and ribs weaken on body chamber which resemblesinner whorls of Pedioceras Gerhardt, 1897 (Crioceratitinae)’(Wright et al., 1983, p. 342).

OccurrenceThe type species and Codazziceras fina Etayo-Serna, 1979

(p. 84, pl. 13, fig. 17, text-fig. 8M,N,Q,S) occur in the LowerConiacian of Colombia. Yubariceras gosavicum Wiedmann,1979 (p. 46, pl. 6, figs C,D) from the Coniacian ofBrandenberg (Tirol, Austria) may also be a Codazziceras.The present record extends the geographic range tonorthern KwaZulu-Natal.

Codazziceras africanum sp. nov., Fig. 1

Derivation of nameAfricanum: of Africa.

TypeThe holotype is SAM-PCZ 022424 (formerly SAS Z1543),

from the Middle or Upper Coniacian part of the St LuciaFormation close to locality 86 of Kennedy & Klinger,(1975), on the southwestern shores of False Bay, LakeSt Lucia, northern KwaZulu-Natal, coordinates28°01’45”S, 32°33’03”E approximately.

DiagnosisA species of Codazziceras with inner umbilical bullae,

inner and outer ventrolateral and siphonal tubercles onprimary ribs alternating regularly with single shorterintercalated ribs with outer ventrolateral and siphonalclavi only.

Dimensions

D Wb Wh Wb:Wh U

SAM-PCZ 022424(ex Z1543)

At: 51.6(100)

– (–) 20.2(38.8)

– 14.9(28.9)

ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 1–2 1

A new species of the Coniacian ammonite genus Codazziceras Etayo-Serna, 1979, previously known with certainty only from Colombia,is described from the St Lucia Formation of northern KwaZulu-Natal.

Keywords: Cretaceous, Coniacian, ammonite, Codazziceras, KwaZulu-Natal, South Africa.

‡In current geopolitical terminology Zululand and Pondoland now form parts of theprovinces of KwaZulu-Natal and the Eastern Cape respectively. For the sake of continu-ity we retain the names Zululand and Natal in the title of our series of systematic descrip-tions of the invertebrate faunas from these regions from 1975 onwards.

*Authors for correspondenceE-mail: [email protected] / [email protected] & [email protected]

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DescriptionThe specimen retains well-preserved iridescent nacreous

shell. Coiling is moderately evolute, with a moderatelywide, shallow umbilicus that comprises 29% of the diameter.The whorl section is slightly compressed in intercostalsection, and only slightly wider than high in costalsection, with the maximum breadth at the umbilicalbullae. The umbilical wall is low and rounded, the flanksflattened and subparallel, with a narrow, flattened venterand rounded ventrolateral shoulder in intercostal section.The costal whorl section is octagonal. There are 12 small,sharp umbilical bullae on the outer whorl. They give riseto a single rib or a pair of distant, straight, narrow,prorsiradiate rounded primary ribs that bear small conicalinner ventrolateral tubercles and outer ventrolateral clavi.Single intercalated ribs arise on the ventrolateral shoulderand bear outer ventrolateral clavi only. All ribs cross theventer and bear elongated siphonal clavi. One primary ribbifurcates at the inner ventrolateral tubercle. There are anestimated 28 ribs at the ventrolateral shoulder of the outerwhorl. The sutures are not exposed.

DiscussionThe presence of alternately long and short ribs, the

former with umbilical, inner and outer ventrolateral andsiphonal clavi, the latter with outer ventrolateral andsiphonal clavi only, distinguish the species. Codazzicerasospinae (Karsten, 1858) (p. 110, pl. 4, fig. 3), as revised byWright et al. 1983 (p. 343, figs 1–4) has inner and outerumbilical tubercles at the same diameter, and the interca-lated ribs bear inner and outer ventrolateral tubercles.Codazziceras fina Etayo-Serna, 1979 (p. 84, pl. 123, fig. 17;text-figs 8M,N,Q,S) has an estimated 34–35 crowded ribsthat are finer than those of the holotype of africanum, theribs frequently arising in pairs, non-bullate primaries, andall ribs have inner and outer ventrolateral tubercles.Yubariceras gosavicum Wiedmann, 1979 (p. 46, pl. 6, figs C,D) may be a Codazziceras. It differs from the present speciesin having inner and outer umbilical tubercles on theprimary ribs, inner and outer ventrolateral tubercles onboth primary and intercalated ribs, and an incipient toweak outer lateral tubercle on some of the ribs.

OccurrenceAs for type.

Kennedy acknowledges the support of the staff of the Geological Collections,Oxford University Museum of Natural History, and the Department of EarthSciences, Oxford, and the financial assistance of the Oppenheimer Fund (Oxford).Klinger acknowledges the support of the staff of the Natural History Collec-tions Department, Iziko South African Museum and financial support from theNRF (South Africa). Thanks are due to referees for their comments on the manu-script.

ABBREVIATIONSD diameter (mm)Wb whorl breadth (mm)Wh whorl height (mm)U umbilicus (mm)

REFERENCESCOOPER, M.R. 1978. Uppermost Cenomanian-basal Turonian ammo-

nites from Salinas, Angola. Annals of the South African Museum 75, 51–152.ETAYO-SERNA, F. 1979. Zonation of the Cretaceous of central Columbia

by Ammonites. Publicaciones Geológicas Especiales del Ingeominas,Bogota 2, 1–186.

GERHARDT, K. 1897. Beiträge zur Kenntniss der Kreideformation inColumbien. Neues Jahrbuch für Mineralogie, Geologie und PaläontologieBeilage Band 11, 118–208.

GROSSOUVRE, A. de 1894. Recherches sur la Craie supérieure, 2,Paléontologie. Les ammonites de la craie supérieure. Mémoires duService de la Carte Géologique détaillée de la France, 1–264 (misdated 1893).Paris, Impremerie Nationale.

HYATT, A. 1889. Genesis of the Arietidae. Smithsonian Contributions toKnowledge 673, xi + 1–239.

KARSTEN, H. 1858. Über die geognostischen Verhältnisse deswestlichen Columbien, der heutigen Republiken Neu-Granada undEquador. Amtlicher Bericht über die Versammlung Deutscher Naturforscherund Ärzte zu Wien (for 1856), 80–117.

KENNEDY, W.J. & KLINGER, H.C. 1975. Cretaceous faunas fromZululand and Natal, South Africa. Introduction, stratigraphy. Bulletinof the British Museum (Natural History) Geology 25, 263–315.

RIEDEL, L. 1938. Amonitas del Cretacico inferior de la CordilleraOriental. In: Estudios geológicos y paleontológicos sobre la CordilleraOriental de Colombia Bogóta 2, 7–78. Bogota, Departamento de Minas yPetroleos, Ministerio de Industrias y Trabajo.

WIEDMANN, J. 1979. Ammonites. Pp. 41–49, In: Herm, D., KAUFFMAN,E.G. & Wiedmann, J. The age and depositional environment of the‘Gosau’-Group (Coniacian-Santonian), Brandenberg/Tirol, Austria.Mitteilungen der Bayerischen Staatssammlung für Paläontologie undhistorische Geologie 19, 27–92.

WRIGHT, C.W., KENNEDY, W.J. & CHANCELLOR, G.R. 1983. Theaffinities of Codazziceras Etayo-Serna, 1979 (Cretaceous Ammonoidea).Cretaceous Research 4, 341–348.

2 ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 1–2

Figure 1. A–D, Codazziceras africanum sp. nov., the holotype, SAM-PCZ 022424 (formerly SAS Z1543), from the Middle or Upper Coniacian part of theSt Lucia Formation close to locality 86 of Kennedy & Klinger (1975) on the southwestern shores of False Bay, Lake St Lucia, northern KwaZulu-Natal,coordinates 28°01’45”S, 32°33’03”E approximately. Figures are ×1.

A C DB

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The ammonite genus Diaziceras Spath, 1921, from theCampanian of KwaZulu-Natal, South Africa,

and MadagascarWilliam James Kennedy1* & Herbert Christian Klinger2*

1Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PW, andDepartment of Earth Sciences, South Parks Road, Oxford OX1 3AN, United Kingdom

2Natural History Collections Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa

Received 23 May 2012. Accepted 18 October 2012

INTRODUCTIONL.F. Spath introduced the genus Diaziceras on the basis

of a single specimen of the type species, Diazicerastissotiaeforme Spath, 1921 (p. 245, pl. 19, figs 1a–k), collectedby A. L. du Toit at Umkwelane Hill, south of Mtubatuba inwhat is now KwaZulu-Natal.

The holotype (Figs 1C–E, 2A,B, 3, 4F) remains the onlyspecimen of the genus known from South Africa, thesystematic position of which has been a matter of debateby subsequent authors. The age of the type specimen isalso uncertain. The type locality corresponds to locality 10of Kennedy & Klinger (1975, p. 282). Here, the basalshell-pebble bed of the St Lucia Formation rests uncon-formably on deeply weathered basalt of the LebomboVolcanics . Concretions three metres above the base haveyielded Placenticeras kaffrarium Etheridge, 1904, andForresteria (Forresteria) alluaudi (Boule, Lemoine &Thévenin, 1907), indicating a Coniacian date for the baseof the sequence (Coniacian II of Kennedy & Klinger(1975)). Taxa recorded from this locality by Spath (1921)also demonstrate the presence of Santonian: i.e.Pseudo-schloenbachia umbulazi (Baily,1855), and Lower Campanian:i.e. Submortoniceras woodsi Spath, 1921. The Parapachydiscussp. nov. aff. colligatus Binkhorst of Spath (1921, p. 226,pl. 22, fig. 1a,b) is the holotype of Menuites (Menuites) spathi(Venzo, 1936) (Kennedy & Klinger, 2006, p. 71, figs 52–68),indicating the presence of an even higher horizon, equiv-alent to Campanian III of Kennedy & Klinger (1975).

It is argued below, on the basis of records of Diazicerasfrom Madagascar that the type specimen is from theLower Campanian. It is also argued that SkoumaliaSummesberger, 1979, from the Santonian of Austria andsouthern France is a synonym of Diaziceras, and that thegenus is closely allied to Eulophoceras, Hyatt, 1903, and amember of the Subfamily Lenticeratinae Hyatt, 1900.

SYSTEMATIC PALAEONTOLOGY

Superfamily Acanthoceratoidea de Grossouvre, 1894Family Sphenodiscidae Hyatt, 1900(= Libycoceratinae Zaborski, 1982, p. 306)Subfamily Lenticeratinae Hyatt, 1900(= Eulophoceratinae Hyatt, 1903;= Diaziceratinae Basse, 1947)

Genus Diaziceras Spath, 1921(= Skoumalia Summesberger, 1979, p. 146)

Type speciesDiaziceras tissotiaeforme Spath, 1921, p. 245, pl. 19,

figs 1a–k, by original designation.

DiagnosisCompressed to moderately inflated, involute. Whorl

section compressed polygonal, with fastigiate venter andstrong siphonal keel. Costal section markedly concavebetween umbilical and ventral tubercles, and on eitherside of strong siphonal keel. Weak to strong umbilicalbullae give rise to pairs of low, broad, prosrsiradiate ribswhich, together with additional intercalated ribs, all bearconical to bullate ventral tubercles. Suture moderately todeeply incised, with asymmetrically bifid to asymmetri-cally trifid E/A and bifid A.

DiscussionThe holotype of the type species is a stout individual

(Figs 1C–E, 2A,B). The holotype of Diaziceras guillantoniHourcq, 1949, p. 22 (108), pl. 12(2), fig. 1; Fig. 9D herein) iscompressed, with weaker ornament of the same basicpattern: umbilical bullae that give rise to pairs of ribs onthe phragmocone, with additional intercalated ribs, Allribs bear ventral tubercles, but these are elongated andprojected forwards, rather than being conical, as inD. tissotiaeforme. The venter is markedly concave on eitherside of the siphonal keel. E/A is asymmetrically trifid,

ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 3–13 3

The ammonite genus Diaziceras Spath, 1921, and the type species, D. tissotiaeforme are revised and referred to the subfamilyLenticeratinae Hyatt, 1900, of the family Sphenodiscidae Hyatt, 1900. Skoumalia Summesberger, 1979, is interpreted as a junior synonymof Diaziceras. Diaziceras guillantoni Hourcq, 1949, and D. spathi Hourcq, 1949, are regarded as synonyms of D. tissotiaeforme, and all arereferred to the Lower Campanian on the basis of records from Madagascar.

Keywords: ammonites, Diaziceras, Campanian, Cretaceous, KwaZulu, South Africa, Madagascar.

*Authors for correspondence.E-mail: [email protected],ac.uk / [email protected] & [email protected]

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and A bifid (Fig. 4B). The differences in ornament betweentissotiaeforme and guillantoni reflect the commoncovariance between whorl inflation and strength orna-ment; they are clearly congeneric.

Skoumalia Summesberger, 1979, with type speciesSkoumalia austriaca Summesberger, 1979 (p. 141, text-figs 26–30; pl. 9, figs 37–41, text-figs 26–30), from the UpperSantonian of the Sandkalkbank of the Bibereckschichtenof the Gosau Basin, Austria, has been regarded as asynonym of Eulophoceras Hyatt, 1903 (Kennedy (1987,p. 776; Kennedy in Kennedy et al. 1995, p. 425; Wright 1996,p. 204). Summesberger recognised two morphotypes in

Skoumalia austriaca. His Form A, which included theholotype (Summesberger, 1979, pl. 9, figs 37, 38; text-figs.26–28; Fig. 4A,C; 9A–C herein) has a compressedphragmocone with fastigiate venter, strong umbilicalbullae that give rise to pairs of straight prorsiradiate ribsthat, together with intercalated ribs, terminate in smallventral bullae. The ornament differs in no significantrespects from that of Diaziceras guillantoni (compareFigs 9A–C and D). The suture of the holotype lacks theventralward part of E/A (Fig. 4A) which is broad, asym-metric, with narrow deep median incision, a deep narrow A,and, bifid A/U2. The suture of a second specimen referred

4 ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 3–13

Figure 1. A, B, the holotype of Diaziceras spathi Hourcq, 1949, p. 107 (21), pl. 12 (2), fig. 2, from the ‘region de Berere’, Madagascar, an unregisteredspecimen in the collections of the École des Mines, Paris, now at the collections of the Université Claude Bernard, Lyon. No precise horizon was given.C–E, Diaziceras tissotiaeforme Spath, 1921, the holotype, by monotypy, SAM-PCZ19040 (formerly 5478 in Spath 1921), the original of Spath 1921, p. 245,pl. 19, fig. 1a–k, from the St Lucia Formation of Umkwelane Hill, northern KwaZulu-Natal. Figures are ×1.

A

C D

B

E

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to form A by Summesberger (1979, text-fig. 28; Fig. 4Cherein) has a broad asymmetric bifid E/A with a deepnarrow incision. The suture is thus less deeply incisedthan in the holotype of Diaziceras tissotiaeforme (Fig. 4F),and lacks the complex folioles adjacent to E, which isobliquely trifid. The bifid E/A and lesser depth of incisionsof Skoumalia austriaca as opposed to the trifid E/A and deepincisions in Diaziceras tissotiaeforme are the only cleardistinguishing feature between the two. The sutures of

other Diaziceras, e.g. D. guillantoni (Fig. 4B) are much lessdeeply incised than in the holotype of D. tissotiaeforme,and that of D. menabense (Fig. 4E) is bifid. Skoumalia isregarded here as a synonym of Diaziceras, rather thanEulophoceras. Summesberger recorded what he regardedas a second form, form B, of Skoumalia austriaca, thatco-occurred with the lectotype (his form A). Form B ofSummesberger (1979, p. 143, text-figs 29, 30; pl. 9,figs 39–41; text-figs 29, 30; 1980, p. 280, pl. 2, figs 5–6; pl. 3,

ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 3–13 5

Figure 2. A, B, Diaziceras tissotiaeforme Spath, 1921, the holotype, by monotypy, SAM-PCZ19040 (formerly 5478 in Spath 1921), the original of Spath(1921, p. 245, pl. 19, fig. 1a–k), from the St Lucia Formation of Umkwelane Hill, northern KwaZulu-Natal. C, D, the holotype of Diaziceras menabenseHourcq, 1949,( p. 109 (23), pl. 13 (3)), fig. 3, from the ‘région de Berere, sommet du Santonien’, Madagascar, an unregistered specimen in the collectionsof the École des Mines, Paris, now housed at the Université Claude Bernard, Lyon. Figures are ×1.

A

C D

B

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6 ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 3–13

figs 7–8; text-figs 5,6) is ornamented by delicate umbilicalbullae on the phragmocone. The outer whorl bears delicatefeebly flexuous growth lines and striae, with ten distantouter lateral bullae. Whereas form A has the ornament ofDiaziceras, Form B (a juvenile from southwest France isshown in Fig. 9E–G for comparison) has delicate ribs andouter lateral bullae, recalling Eulophoceras, as discussedelsewhere (Kennedy & Klinger, 2012, in press).

Spath (1921, p. 242) was unequivocal in his view of theaffinities of Diaziceras: ‘Its suture-line stamps it as beingnear to the genera Eulophoceras Hyatt and Spheniscoceras’Spath also noted the similarity in ornament betweenDiaziceras and certain Tissotidae Hyatt, 1900, specificallyMetatissotia fourneli (Bayle, 1878) ( pl. 40, fig. 3), butdismissed the possibility of affinity, given the pseudo-ceratitic suture of the family. In contrast, Hourcq (1949,p. 116 (30)), placed Diaziceras and Eulophoceras in thefamily Tissotidae Hyatt, 1900. Basse (1947, p. 159 (63))introduced a subfamily Diaziceratinae, which she placedin the Tissotidae. Wright (1957, p. L437) assigned the genusto the Lenticeratinae Hyatt, 1900 (= EulophoceratinaeHyatt, 1903). In 1996 (p. 189) he preferred placement in thesubfamily Barroisiceratinae Basse, 1947, of the familyCollignoniceratidae Wright & Wright, 1951. An orphangenus indeed!

We revert to the views of Spath. The sutures of Diazicerasand Eulophoceras share an asymmetrically trifid E/A.The weak ribs with no or feeble umbilical bullae, andstrengthened rib terminations (but only rarely bullae) ofEulophoceras are a reflection of the compressed, oxyconewhorl section. Diaziceras tissotiaeforme has inflated whorlsand strong ribs and tubercles. The ribs and tubercles of

compressed Diaziceras guillantoni Hourcq, 1949 (p. 108(22), pl. 12 (2), fig. 1; text-fig. 15) are intermediate, differ-ences that reflect covariance between ornament strengthand whorl section.

OccurrenceUpper Santonian of the Gosau Basin, Austria, and

Corbières in southern France. Lower Campanian ofMadagascar (following Collignon, 1969, although origi-nally referred to the top of the Santonian by Hourq, 1949).On the basis of the Madagascan occurrences, the typeoccurrence in KwaZulu-Natal is inferred to be in theLower Campanian.

Diaziceras tissotiaeforme Spath, 1921Figs 1C–E, 2–3, 4E,F, 5

1921 Diaziceras tissotiaeforme Spath, p. 245, pl. 19, figs 1a–k.1949 Diaziceras Spathi Hourcq, p. 107 (21), pl. 12 (2), fig. 2.1949 Diaziceras menabense Hourcq, p. 109 (23), pl. 13 (3), fig. 3;

text-figs 16, 17.1957 Diaziceras tissotiaeforme Spath; Wright, p. L437, text-

fig. 552, 4a–c.1969 Diaziceras spathi Hourcq; Collignon, p. 210, pl. 603,

fig. 2258.1996 Diaziceras tissotiaeforme Spath; Wright, p. 189, text-fig. 143,

2a–c.

TypeThe holotype, by monotypy, is SAM-PCZ19040

(formerly 5478 in Spath 1921), the original of Spath 1921,p. 245, pl. 19, figs 1a–k, from the St Lucia Formation ofUmkwelane Hill, northern KwaZulu-Natal, locality 10 ofKennedy & Klinger (1975, p. 282).

Figure 3. Diaziceras tissotiaeforme Spath, 1921, the holotype, by monotypy, SAM-PCZ19040 (formerly 5478 in Spath 1921), the original of Spath (1921,p. 245, pl. 19, fig. 1a–k), from the St Lucia Formation of Umkwelane Hill, northern KwaZulu-Natal. A, the inner whorls at a diameter of 3 mm; copy ofSpath, (1921, pl. 1, fig.1d); B, development of the suture line at i, a diameter of 1.5 mm; ii at 2 mm; iii at 3 mm; iv at 5.5 mm; v at 8 mm. Copy of Spath,(1921, pl. 1, fig. 1g–1k). Scale bars are 1 mm.

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Dimensions of the holotype

D Wb Wh Wb:Wh U

PCZ19040,c 80.6 (100) 47.7 (59) 42.1 (52) 1.13 13.6 (16)

PCZ19040,ic 80.6 (100) 38.6 (47) 42.1 (52) 0.92 13.6 (16)

DescriptionThe holotype is a well-preserved undeformed individual

retaining traces of the original aragonitic shell. It is septateto a diameter of 81.4 mm, the last few septa crowded,

suggesting the specimen may be the phragmocone of anadult. A short, worn section of body chamber survives,and extends the specimen to a maximum preserveddiameter of 84.5 mm. Coiling is very involute, with a small,very deep conical umbilicus that comprises 16.9% of thediameter, the umbilical wall flattened and outwardly-inclined. The umbilical shoulder is broadly rounded. Theintercostal whorl section is compressed polygonal, withthe greatest breadth just outside the umbilical shoulderand a whorl breadth to height ratio of 0.92. The innerflanks are broadly rounded, the outer flanks flattened and

Figure 4. Suture lines. A, Skoumalia austriaca Summesberger, 1979, form A (copy of Summesberger 1979, text-fig. 27). B, the holotype of Diazicerasguillantoni Hourcq, 1949, p. 108 (22), pl. 12 (2), fig. 1. C, Skoumalia austriaca Summesberger, 1979, form A, (copy of Summesberger 1979, text-fig. 28.D, Skoumalia austriaca Summesberger, 1979, form B (copy of Summesberger 1979, text-fig. 30). E, the holotype of Diaziceras menabense Hourcq, 1949,p. 109 (23), pl. 13 (3), fig. 3. F, the holotype of Diaziceras tissotiaeforme Spath, 1921, copy of Spath (1921, pl. 19, fig. 1e, f.). Scale bar is 10 mm.

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Figure 5. Diaziceras tissotiaeforme Spath, 1921, the original of Diaziceras spathi Hourcq, 1941, sensu Collignon, (1969, pl. 603, fig. 2258), from the ‘base duCampanien Inférieur. Zone à Anapachydiscus wittekindi et Eulophoceras jacobi. Sous-Zone à Besairiella besairiei (base). Gisement 195 de la Coupe deBerere II-B (Belo-sur-Tsiribihina)’, Madagascar. The figure is ×1.

Figure 6. External suture of Diaziceras guillantoni Hourcq, 1949, the original of Collignon, (1969, pl. 614, fig. 2261). Scale bar is 10 mm.

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convergent, the venter obtusely fastigiate, with a strongsiphonal keel. The greatest breadth is at the massiveumbilical bullae in costal section, with a whorl breadth toheight ratio of 1.13. The whorl section is concave betweenumbilical bullae and ventrolateral tubercles, and betweenventrolateral tubercles and keeled venter. There are fivemassive umbilical bullae, perched on the umbilical shoulder.The bullae give rise to pairs of low, broad, convexprorsiradiate ribs that sweep across the flanks and link tosmall conical ventral tubercles. One or more short ribsintercalate between the successive pairs of bullate prima-ries, to give a total of 12 ribs on the adapertural half of theouter whorl, and an estimated 24 per whorl.

The mature suture (Fig. 4F) has a broad, deeply incisedasymmetrically trifid E/A, A with deep incisions and asmall, bifid U2. The early sutural development is shown inFig. 3.

DiscussionDiaziceras menabense Hourcq, 1949 (p. 23 (109),

text-figs 16,17; pl. 13 (3), fig. 3) is a synonym ofD. tissotiaeforme in our view. The holotype (Figs 2C,D, 4E)is an unregistered specimen in the collections of the Écoledes Mines, Paris, now housed at the Université Claude

Bernard, Lyon. It is from Berere, Madagascar, and wasreferred to the top of the Santonian by Hourcq. A wornand battered internal mould with a 60° sector of bodychamber preserved, the dimensions are a follows:

D ic: 98.0 (100) Wb: 37.5 (38.3) Wh: 54.0 (55.1) Wb:Wh:0.69 U:10.7 (10.9)

Coiling is very involute, the small deep umbilicuscomprising 10.9% of the diameter, with a flattened, out-ward-inclined wall and a broadly rounded umbilicalshoulder. The intercostal whorl section is compressedlanceolate, with the greatest breadth just outside theumbilical shoulder. Nine to 10 conical umbilical tuberclesperch on the umbilical shoulder; strong on the phragmo-cone, they weaken markedly on the body chamber. Onthe phragmocone, the bullae give rise to pairs of lowbroad prorsiradiate ribs that terminate in conical outerlateral tubercles. Beyond the tubercles the flanks areconcave and converge to the acute venter. The tuberclesand flank ribs persist onto the body chamber, but theventer becomes less acute. The suture (Fig. 4E) has abroad, little-incised incipiently obliquely trifid E/A, anarrow A, and little-incised bifid U2. The ornament isslightly weaker, and the whorl section compressed whencompared to that of the holotype of tissotiaeforme while the

Figure 7. Diaziceras guillantoni Hourcq, 1949, the original of Collignon, (1969, pl. 604, fig. 2261), from ‘base du Campanien Inférieur. Zone àAnapachydiscus wittekindi et Eulophoceras jacobi. Sous-Zone à Hourcquiella hourcqui. Gisement 195 de la Coupe de Berere II-B (Belo-sur-Tsiribihina)’,Madagascar. The figure is ×1.

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suture is much less deeply incised, lacking the longcomplx folioles on E/A adjacent to A, and barely trifid.

Diaziceras spathi Hourcq, 1949 (p. 107 (21), pl. 12 (2), fig. 2)is a further synonym. The holotype, by monotypy(Fig. 1A,B), is an unregistered specimen in the collectionsof the École des Mines, Paris, now housed at the UniversitéClaude Bernard, Lyon. It from Berere, Madagascar. Aworn, wholly septate internal mould, the dimensions areas follows:

D ic: 88.5 (100); Wb: 42.0 (0.47); Wh:47.5 (0.54) Wb:Wh0.89 U:15.5 (17.5)

Coiling is involute, the deep umbilicus comprising 17.5%of the diameter, with a flattened, outward-inclined umbil-ical wall and broadly rounded umbilical shoulder. Thewhorl section is stoutly lanceolate in intercostal section,with the greatest breadth just outside the umbilical shoulder.The greatest breadth is at the umbilical bullae in costal

section. These number eight per whorl, and are perchedon the umbilical shoulder. They give rise to pairs of low,broad blunt straight prorsiradiate ribs that terminate inconical outer lateral tubercles, 19–20 per whorl. The flanksare concave in costal section between these tuberclesand the obtuse venter. The specimen has a slightly morecompressed whorl section than the holotype of D. tisso-tiaeforme, but is otherwise identical in our view (compareFigs 1A,B and 1C–E).

Diaziceras guillantoni Hourcq, 1949 (p. 108 (22), pl. 12 (2),fig. 1; text-fig. 15), is a distinct compressed form (Fig. 9D).The holotype by monotypy is an unregistered specimenin the collections of the École des Mines, Paris, nowhoused at the Université Claude Bernard, Lyon. It is fromBerere, Madagascar, and was referred to the top of theSantonian by Hourcq. A worn internal mould with a 180°sector of body chamber preserved, and a near-complete

Figure 8. Diaziceras guillantoni Hourcq, 1949, the original of Collignon, (1969, pl. 604, fig. 2261), with the previously unfigured body chamber fragmentincluded, from the ‘base du Campanien Inférieur. Zone à Anapachydiscus wittekindi et Eulophoceras jacobi. Sous-Zone à Hourcquiella hourcqui. Gisement195 de la Coupe de Berere II-B (Belo-sur-Tsiribihina)’, Madagascar. The figure is ×1.

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Figure 9. A–C, the holotype of Skoumalia austriaca Summesberger, 1979, SK,N.1977/14, the original of Summesberger, (1979, pl. 9, figs 37, 38) from theUpper Santonian Sandkalkbank of the Bibereckschichten of the Gosau Basin, Upper Austria. D, the holotype of Diaziceras guillantoni Hourcq, 1949,the original of Hourcq, (1949, pl. 12 (2), fig. 1), from the ‘Région de Berere, sommet du Santonien.’ E–G, Skoumalia austriaca Summesberger, 1979,form B of Summesberger, (1980), A363, from the Upper Santonian of Autoroute cutting S2, west of Saintes, Charente-Maritime, France in the collec-tions of the Université Claude Bernard. All figures are ×1.

A

C

D

B

E

F G

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adult, the dimensions of which are as follows:

D: 132.0 (100) Wb: 40.5 (30.7) Wh: 70.0 (53.0) Wb:Wh0.58 U:18.0 (13.6)

Coiling is involute, the umbilicus small and deep, com-prising 13.6% of the diameter, with an outward-inclinedwall and broadly rounded umbilical shoulder. The whorlsection is compressed lanceolate, with a whorl breadth toheight ratio of 0.58. Four massive bullae perch on theumbilical shoulder of the adapical half of the outer whorl.They give rise to pairs of low, broad, straight prorsiradiateribs with occasional intercalated ribs arising aroundmid-flank, to give a total of ten ribs per half whorl on theouter flank, where the ribs strengthen into conical tofeebly bullate tubercles. The umbilical bullae efface pro-gressively on the body chamber and the flank ribsweaken. The outer lateral tubercles become markedlybullate and prorsiradiate before weakening at the greatestpreserved diameter. The suture (Fig. 4B) has a very broad,obliquely trifid E/A, with deep incisions and plumpfolioles; A is asymmetrically bifid; U2 is broad, bifid withonly minor incisions. It is interpreted as a microconch. Themacroconch is represented by the original of Collignon(1969, pl. 604, fig. 2261, see Figs 6–8 herein) from the LowerCampanian Hourcquiella bererensis Subzone of Collignon’s‘Zone à Anapachydiscus wittekindi et Eulophoceras jacobi’ ofBerere (Belo-sur-Tsiribihina), Madagascar. The umbilicalbullae are weak and very elongate, the ribs straight andprorsiradiate, arising from the bullae in pairs and interca-lating, with an oblique prorsiradiate ventrolateral bulla onthe phragmocone. Ribs and tubercles weaken and effaceon the body chamber, which occupies a 180° sector andmay be incomplete (Fig. 8). The sutures (Fig. 6) arecrowded and interfere. The saddles are broad and plump,with A narrower.

Diaziceras austriaca (Summesberger, 1979), is discussedabove, and figured here as Figs 4A,C, 9A–C, E–G. It com-bines a compressed whorl section with massive umbilicaltubercles and weak flank ribs in form A of Summesberger,the suture (Figs 4A,C) is a little less deeply incised thanthat of D. tissotiaeforme, and has E/A obliquely asymmetri-cally bifid, rather than trifid.

The holotype of Diaziceras tissotiaeforme is interpreted asa microconch. Collignon (1969, pl. 603, fig. 2258) figured(as D. spathi) a worn and battered individual 150 mm indiameter with a 180° sector of body chamber preservedthat is interpreted as a near-complete macroconch (Fig. 5).It has seven massive bullae perched on the umbilicalshoulder that give rise to pairs of ribs on the phragmocone,the ribs on the body chamber simple, to give a total of17 ribs on the outer whorl that terminate in strongventrolateral tubercles. It is from the base of theCampanian, the base of the Besairiella besairiei Subzone ofCollignon’s Zone à Anapachydiscus wittekindi et Eulophocerasjacobi of Berere (Belo-sur-Tsiribihina), Madagascar.

OccurrenceThe type occurrence at Umkwelane Hill in KwaZulu-

Natal is imprecisely dated, but inferred to be LowerCampanian on the basis of the record of Collignon (1969)from Madagascar.

Kennedy acknowledges the support of the staff of the Geological Collections,Oxford University Museum of Natural History, and the Department of EarthSciences, Oxford, and the financial assistance of the Oppenheimer Fund (Oxford).Klinger acknowledges financial support from the NRF (South Africa) and helpfrom the staff of the Natural History Collections Department, Iziko, South AfricanMuseum. Thanks are due to the reviewers for constructive comments on themanuscript.

ABBREVIATIONSDimensions are given in millimetres.D diameterWb whorl breadthWh whorl heightU umbilicusc costal dimensionic intercostal dimensionFigures in brackets are dimensions as a percentage of the diameter.

The suture terminology is that of Korn et al. (2003):E external lobeA adventive lobe (= lateral lobe, L, of Kullmann & Wiedmann 1970)U umbilical lobeI internal lobe

REFERENCESBAILY, W.H. 1855. Description of some Cretaceous fossils from South

Africa. Quarterly Journal of the Geological Society of London 11, 454–465.BASSE, E. 1947. Les peuplements Malgaches de Barroisiceras (Révision

du genre Barroisiceras de Gross). Paléontologie de Madagascar 26.Annales de Paléontologie 22, 97–190.

BAYLE, É. 1878. Fossiles principaux des terrains. Explication de la CarteGéologique de France 4(1) (Atlas), 158 pls. Paris, Service de la CarteGéologique detaillée.

BOULE, M., LEMOINE, P. and THÉVENIN, A. 1906–1907. Paléontologiede Madagascar III Céphalopodes crétacés des environs deDiego-Suarez. Annales de Paléontologie 1, 173–192 (1–20) (1906) 2, 1–56(21–76) (1907).

COLLIGNON, M. 1969. Atlas des fossiles caractéristiques de Madagascar(Ammonites). XV, (Campanien inférieur), xi + 1–216. Tananarive: ServiceGéologique.

ETHERIDGE, R. 1904. Cretaceous fossils of Natal. 1. The Umkwelane HillDeposit. Report of the Geological Survey of Natal and Zululand 1, 71–93.

GROSSOUVRE, A. de 1894. Recherches sur la craie supérieure, 2,Paléontologie. Les ammonites de la craie supérieure. Mémoires duService de la Carte Géologique détaillée de la France 1, 1–264 (misdated1893). Paris, Imprimerie national .

HOURCQ, V. 1949. Paléontologie de Madagascar. XXVIII. Sur quelquesammonites du Sénonien. Annales de Paléontologie 35, 10(87) – 31(117).

HYATT, A. 1900. Cephalopoda. In: Zittel, K.A. von 1896–1900, Textbook ofPalaeontology (transl. Eastman, C.R.), 502–604. London and New York,Macmillan.

HYATT, A. 1903. Pseudoceratites of the Cretaceous. United States Geologi-cal Survey Monograph 44, 351 pp.

KENNEDY, W.J. 1987. Ammonites from the type Santonian and adjacentparts of northern Aquitaine, western France. Palaeontology 30, 765–782.

KENNEDY, W.J., BILOTTE, M. & MELCHIOR, P. 1995. Ammonitefaunas, biostratigraphy and sequence stratigraphy of the Coniacian-Santonian of the Corbières. Bulletin des Centres de Recherche Explorationet Production Elf-Aquitaine 19, 377–499.

KENNEDY, W.J. & KLINGER, H.C. 1975. Cretaceous faunas fromZululand and Natal, South Africa. Introduction, Stratigraphy. Bulletinof the British Museum (Natural History) Geology 25, 263–315.

KENNEDY, W.J. & KLINGER, H.C. 2006. Cretaceous faunas fromZululand and Natal, South Africa. The ammonite familyPachydiscidae Spath, 1922. African Natural History 2, 17–168.

KENNEDY, W.J. & KLINGER, H.C. 2012, in press. Cretaceous faunasfrom Zululand and Natal, South Africa. The Santonian-Campanianammonite genus Eulophoceras Hyatt, 1903. African Natural History 8.

KORN, D., EBBIGHAUSEN, V., BOCKWINKEL, J. & KLUG, C. 2003. TheA-mode sutural ontogeny in prolecanitid ammonoids. Palaeontology46, 1123–1132.

KULLMANN, J. & WIEDMANN, J. 1970. Significance of sutures inphylogeny of Ammonoidea. University of Kansas, Paleontological Contri-butions 42, 1–32.

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SPATH, L.F. 1921. On Cretaceous Cephalopoda from Zululand. Annals ofthe South African Museum 12, 217–321.

SUMMESBERGER, H. 1979. Eine obersantone Ammonitenfauna ausdem Becken von Gosau (Oberösterreich). Annalen des Natur-historischen Museums Wien 83, 109–176.

SUMMESBERGER, H. 1980. Neue Ammoniten aus der Sandkalkbankder Hochmoosschichten (Obersanton; Gosau, Austria). Annalen desnaturhistorischen Museums Wien 83, 275–383.

VENZO, S. 1936. Cefalopodi del Cretaceo medio-superiore delloZululand. Palaeontographia Italica 36, 59–133 (1–75).

WRIGHT, C.W. 1957. [Cretaceous Ammonoidea]. In: Moore, R.C. (ed.),Treatise on Invertebrate Paleontology. Part L, Mollusca 4, Cephalopoda

Ammonoidea. Boulder, New York and Lawrence, Geological Society ofAmerica and University of Kansas Press.

WRIGHT, C.W. 1996. In: Treatise on Invertebrate Paleontology. Part L,Mollusca 4: Cretaceous Ammonoidea (with contributions by J.H.Calloman (sic) and M.K. Howarth). Boulder, Colorado and Lawrence,Kansas, Geological Society of America and University of Kansas.

WRIGHT, C.W. & WRIGHT, E.V. 1951. A survey of the fossil Cephalopodaof the Chalk of Great Britain. Monograph of the Palaeontographical SocietyMonographs, London, 140, 1–40.

ZABORSKI, P.M.P. 1982. Campanian and Maastrichtian sphenodiscidammonites from southern Nigeria. Bulletin of the British Museum (Natu-ral History) Geology 36, 303–332.

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Cretaceous faunas from Zululand and Natal,South Africa‡. A new species of the ammonite genus

Salaziceras Breistroffer, 1936, from the LowerCenomanian Mzinene Formation

William James Kennedy1* & Herbert Christian Klinger2*1Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PW, andDepartment of Earth Sciences, South Parks Road, Oxford OX1 3AN, United Kingdom

2Natural History Collections Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa

Received 6 August 2012. Accepted 18 October 2012

INTRODUCTIONWe have previously recorded the type species of the

diminutive Late Albian ammonite genus SalazicerasBreistroffer, 1936, from the Upper Albian part of theMzinene Formation at Ndumu in northern KwaZulu-Natal (Klinger & Kennedy 1994), and interpreted theenigmatic Acanthoceras naviculare from Mont Raynaud,Madagascar, described by Boule et al. (1907, p. 30 (10), pl. 1(8), fig. 1 ) as a hypermorphic giant species of this normallydiminutive genus (Kennedy & Klinger 2008). Revision ofthe KwaZulu-Natal Stoliczkaiinae (in progress), led tothe recognition of a second specimen of this genus fromKwaZulu-Natal, which we describe below, and refer to anew species, Salaziceras simplex.

SYSTEMATIC PALAEONTOLOGY

Superfamily Acanthoceratoidea de Grossouvre, 1894Family Flickiidae Adkins, 1928Subfamily Salaziceratinae Wright & Kennedy, 1984

Genus Salaziceras Breistroffer, 1936

Type speciesAmmonites salazacensis Hébert & Munier-Chalmas,

1875, p. 114, pl. 5, fig. 6, by original designation byBreistroffer (1936, p. 64).

DiscussionSee Wright & Kennedy (1979, p. 686).

OccurrenceUpper Upper Albian, southern England, southeast

France, Hungary, Morocco, Nigeria, northern KwaZulu-

Natal, South Africa, and Madagascar. Lower Cenomanianof northern KwaZulu-Natal, South Africa.

Salaziceras simplex sp. nov., Fig. 1

Derivation of nameSimplex: simple.

TypeThe holotype is SAM-PCZ022425 (formerly D 2945, ex

E.C.N. Van Hoepen Collection), from the LowerCenomanian Mzinene Formation at the Skoenberg, corre-sponding to locality 61 of Kennedy & Klinger (1975, p. 189,fig. 6), NNW of Hluhluwe, northern KwaZulu-Natal,coordinates 27°52’17”S, 32°20’19”E.

DiagnosisA species of Salaziceras in which the earlier phragmocone

whorls are ornamented by coarse bullate primary ribsseparated by one or two intercalated ribs that extendacross the venter, followed by a late phragmocone andearly body chamber phase ornamented by bullate primaryribs only that efface on the ventrolateral shoulders andventer.

Dimensions

D Wb Wh Wb:Wh U

SAM-PCZ022424

(ex D2945)

16.4 (100) 10.4 (64.0) 9.0 (54.9) 1.17 – (–)

at 24.8 (100) 10.4 (44.0) 10.0 (40.3) 1.1 6.9 (27.8)

DescriptionThe holotype is a well-preserved phragmocone and a

120° sector of body chamber, with extensive areas oflimonitized shell preserved that largely obscure the sutures.Coiling is evolute, the umbilicus comprising 27.8% of thediameter, of moderate depth, with a convex wall andbroadly rounded umbilical shoulder. The whorl section is

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A diminutive ammonite collected by E.C.N. Van Hoepen from the Cenomanian part of the Mzinene Formation of the Skoenberg innorthern KwaZulu-Natal, is described as Salaziceras simplex sp. nov., and interpreted as one of the last survivors of the genus.

Keywords: Cretaceous, Cenomanian, Salaziceras, ammonite, KwaZulu-Natal, South Africa.

‡In current geopolitical terminology Zululand and Pondoland now form parts of theprovinces of KwaZulu-Natal and the Eastern Cape. For the sake of continuity we retainthe names Zululand and Natal in the title of our series of systematic descriptions of theinvertebrate faunas from these regions from 1975 onwards.

*Authors for correspondenceE-mail: [email protected] / [email protected] & [email protected]

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depressed reniform, with the greatest breadth at theumbilical bullae and a costal whorl breadth to height ratioof up to 1.3 on the adapertural parts of the phragmocone.The intercostal whorl breadth to height ratio is 1.1 on thebody chamber at the greatest preserved diameter. Thenucleus of the specimen at 16.4 mm diameter has anestimated 12 progressively strengthening umbilical bullaeper whorl. These give rise to low, broad, prorsiradiate ribsor, occasionally, a pair of ribs, while one or two shorter ribsintercalate low on the flank. The ribs strengthen across theouter flank before weakening, and crossing the venter in avery feeble convexity. Towards the end of the nucleus theribs are near effaced on the venter. On a 120° sector at theadapertural end of the phragmocone and adapical end ofthe body chamber four strong, distant umbilical bullaegive rise to a single prorsiradiate rib that effaces on theventrolateral shoulders and venter, which are nearsmooth. The final 60° sector bears a single non-bullate rib,and is thereafter smooth; this change in ornament sug-gesting the proximity of the adult aperture. The suturesare not seen.

DiscussionSalaziceras simplex differs from typical forms of the type

species, S. salazacense salazacense in the absence of ribs onthe ventrolateral shoulders and venter of the latest partsof the phragmocone and body chamber (see illustrationsin Scholz,1979, p. 92, pl. 21, figs 6–10, 13–15, 17; text-figs 25,26A,B,H,I,J,L,M,U,V and Szives, 2007, p. 107, pl. 14, fig. 14;pl. 16, figs 7–12; pl. 20, figs 6, 8, 9 and references therein).Salaziceras salazacense gracilicostatus Scholz 1979 (p. 95,pl. 21, figs 11, 12; text-fig. 25) is a slender form, withcrowded ribs that are well developed across the venter

throughout the known ontogeny, and lack umbilicalbullae on the primary ribs. Salaziceras salazacensepeyrolasense Scholz 1979,( p. 93, pl. 21, figs 16, 18–20;text-figs 25, 26B, 27C–G,K,N,O,P,Q) is near smooth but forconstrictions at a diameter that corresponds to theadapertural section of the nucleus of the holotype of sim-plex, thereafter developing very distant flank ribs, threeper half whorl, lacking strong bullae. Salaziceras lemoineiKennedy & Klinger, 2008 (p. 115, text-figs 1G–K, 2) is muchlarger (up to 53 mm in diameter), the primary ribs lackingumbilical bullae, and alternating with short intercalatedribs, all ribs strongly developed on the ventrolateralshoulders and venter of the adult body chamber.

OccurrenceAs for type.

ABBREVIATIONSD diameter (mm)Wb whorl breadth (mm)Wh whorl height (mm)U umbilicus (mm)

Kennedy acknowledges the support of the staff of the Geological Collections,Oxford University Museum of Natural History, and the Department of EarthSciences, Oxford, and the financial assistance of the Oppenheimer Fund (Oxford).Klinger acknowledges the technical support of the staff of the Natural HistoryCollections Department, Iziko, South African Museum and financial support fromthe NRF (South Africa).

REFERENCESADKINS, W.S. 1928. Handbook of Texas Cretaceous fossils. University of

Texas Bulletin 2838, 1–385.BOULE, M., LEMOINE, P. & THÉVENIN, A. 1906–1907. Paléontologie de

Madagascar III Céphalopodes crétacés des environs de Diego-Suarez.Annales de Paléontologie 1, 173–192 (1–20); 2, 1–56 (21–76).

BREISTROFFER, M. 1936. Les subdivisions du Vraconien dans le

16 ISSN 0078-8554 Palaeont. afr. (December 2012) 47: 15–17

Figure 1. Salaziceras simplex sp. nov. The holotype is SAM-PCZ 022425 (formerly D. 2945, ex E.C.N. Van Hoepen Collection), from the LowerCenomanian Mzinene Formation at the Skoenberg, corresponding to locality 61 of Kennedy & Klinger (1975, p. 189, fig. 6), NNW of Hluhluwe,northern KwaZulu-Natal, coordinates 27°52’17’’S, 32°20’19’’E. A–C are ×1; D–L are ×2.

A C

D

B

E F G

H I J K L

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Sud-Est de la France. Bulletin de la Société Géologique de France (6)5,63–68.

GROSSOUVRE, A. de 1894. Recherches sur la craie supérieure, 2,Paléontologie. Les ammonites de la craie supérieure. Mémoires duService de la Carte Géologique détaillée de la France. 264 pp.(misdated1893). Paris, Imprimerie Nationale.

HÉBERT, [E.] & MUNIER-CHALMAS, [E.P.A.]. 1875. Fossiles du Bassind’Uchaux. Annales des Sciences Géologiques, Paris 6, 113–132.

KENNEDY, W.J. & KLINGER, H.C. 1975. Cretaceous faunas fromZululand and Natal, South Africa. Introduction, Stratigraphy. Bulletinof the British Museum (Natural History) Geology 25, 263–315.

KENNEDY, W.J. & KLINGER, H.C. 2008. Hypermorphosis in Salaziceras,a Cretaceous ammonite, from Madagascar. African Natural History 4,113–116.

KLINGER, H.C. & KENNEDY, W.J. 1994. Cretaceous faunas from

Zululand and Natal. Salaziceras salazacense Hébert & Munier-Chalmas,1875) from the Mzinene Formation of northern Zululand. SouthAfrican Journal of Geology 97(3), 146–148.

SCHOLZ, G. 1979. Die Ammoniten des Vracon (Oberalb, dispar Zone)des Bakony-Gebirges (Westungarn) und eine Revision der wichtigstenVracon-Arten der Westmediterranen Faunenprovinz. Palaeonto-graphica A165, 1–136.

SZIVES, O. 2007. Albian Stage. In: Aptian-Campanian ammonites ofHungary. Geologica Hungarica, Series Palaeontologica 57, 75–122.

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WRIGHT, C.W. & KENNEDY, W.J. 1984. The affinities of the Creta-ceous ammonite Neosaynoceras Breistroffer, 1947. Palaeontology 27,159–167.

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A new caprin bovid (Mammalia) from the late Mioceneof Morocco

Denis Geraads1*, Siham El Boughabi2 & Samir Zouhri2

1CNRS, UPR 2147, 44 rue de l’Amiral Mouchez, F-75014 Paris, France2Laboratoire de Géosciences, Faculté des Sciences, Université Hassan II-Casablanca,

Km 8, route d’El Jadida, BP 5366 Maârif, 20100 Casablanca, Morocco

Received 23 April 2012. Accepted 20 August 2012

INTRODUCTIONWith more than 70 living species, the Bovidae are by far

the most diverse family of large mammals in Africa today.They include representatives of all major tribes except theBoselaphini, but most of them belong to tribes that aretoday endemic to the Afro-Arabian domain, namely theTragelaphini, Cephalophini, Neotragini, Aepycerotini,Reduncini, Hippotragini and Alcelaphini. The Antilopiniare also well represented, but they are mainly a Palearcticgroup. In the Pliocene and Pleistocene, evidence from thewhole continent shows that the same groups dominatedthe bovid assemblages, although the proportions of thevarious tribes vary sharply with age and location. To thesegroups, Ethiopian in the terminology of modern faunalprovinces, we must add the Bovini, more common andmore diverse than today, the Boselaphini that wentextinct in Africa by the earliest Pliocene (their last appear-ance is at Langebaanweg in South Africa, and in the ApakMember of Lothagam in Kenya), and the Caprini thatwere far less common than in Eurasia. It is likely that thedifferentiation of the African tribes took place in theMiocene, but it is only by the late Miocene, c. 7 Ma, atSahabi in Libya, Lothagam in Kenya, and Toros Menallain Chad, that we find fossils that can be unambiguouslyassigned to these modern endemic tribes (Lehmann &Thomas 1987; Harris 2003; Geraads et al. 2008; Bibi et al.2009; Gentry 2010). At that time, the composition of thebovid assemblage is already fully African in character,with the exception of the last boselaphines.

Earlier late Miocene African bovid faunas are poorlyknown. The Beglia Formation of Tunisia, mostly, if notfully (Geraads 1989) of late Miocene age, yielded severalbovids but only a few have been described (Robinson1972, 1986); a few fragmentary specimens were describedby Geraads (1989) from the slightly younger sites of JebelKrechem in the same country; Bou Hanifia in Algeria

yielded only the very poorly known Damalavus boroccoiArambourg, 1959. It is noteworthy that none of the fossilsfrom these localities foreshadow the later endemic Africantribes. The greatest potential for improving our knowl-edge of bovids of the first part of the late Miocene(Vallesian – equivalent) in Africa probably rests in theNakali and Samburu beds of Kenya, but these have notyet been published.

In this paper we describe the most complete knownskull of a North African Miocene bovid, and assign it to anew taxon. It comes from the fluvial deposits (coarsesands and conglomerates) of the Upper Member of the AïtKandoula Formation, near the village of Skoura, East ofOuarzazate, south of the High Atlas of Morocco (Fig. 1);these deposits have been dated to the late Miocene on afaunal basis (Zouhri et al. 2012).

SYSTEMATIC PALEONTOLOGYIn the descriptions, the tooth rows are supposed to be

horizontal. Upper teeth are in upper case, lower teeth arein lower case.

Family Bovidae Gray, 1821Subfamily Antilopinae Gray, 1821Tribe Caprini Gray, 1821

Genus Skouraia, gen. nov.Type species. Skouraia helicoides, new species.Derivatio nominis. From Skoura, the village closest to the

area of the type locality.Diagnosis. That of the type-species.

Skouraia helicoides sp. nov., Fig. 2Holotype. Almost complete cranium, bearing the complete

right horn-core and a large part of the left one, with mostof the teeth except the left P2 and all right premolars, butlacking most of the dorsal part of the face anterior to theorbit, and the premaxillae. It is unnumbered and preserved

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*Author for correspondence. E-mail: [email protected]

We describe here a bovid skull from the Upper Member of the Aït Kandoula Formation near Ouarzazate, Morocco, which can be datedby biostratigraphy to the late Miocene, Turolian-equivalent. We assign it to a new taxon, Skouraia helicoides, gen. nov., sp. nov. It has longhorn-cores that are much inclined backwards, strongly spiralled in homonymous direction, very divergent, and have a stronganterolateral keel. The strong cranial flexure, broad basioccipital, and aegodont teeth demand inclusion of this new taxon within thetribe Caprini, a mostly Eurasian group with few African representatives. Skouraia must be an early offshoot of this tribe, but its highlyderived cranial features suggest that the Caprini may have experienced, in the poorly known late Miocene of Africa, a broader morpho-logical diversification than in Europe.

Keywords: Upper Miocene, Morocco, Africa, Bovidae, Caprinae, Caprini.

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in the private museum of Mr Brahim Tahiri in Erfoud,Morocco.

Diagnosis. A bovid with strong cranial flexure, early fusionof the bones of the braincase, low occipital, basioccipitalbroad and rather flat, with a narrow central groove.Horn-cores strongly divergent, much inclined posteriorly,with a virtually circular cross-section except for a strongantero-lateral keel, and describing a strong homonymousspiral. Upper teeth aegodont, with narrow styles, flat orpoorly convex labial walls, and short premolars.

Type locality. The specimen was collected near TiziN’Tadderht, in the Upper Member of the Aït KandoulaFormation of the Ouarzazate basin, Morocco, associatedwith a fauna dated by biostratigraphy to the youngestpart of the late Miocene (equivalent to the EuropeanTurolian; Zouhri et al. 2012).

Derivatio nominis. Because the horn-cores are stronglyspiralled.

Remarks. It is of course unfortunate that the holotype,and only known specimen, be kept in a private (albeitfreely accessible as of November 2010) museum. Still, webelieve that in spite of this shortcoming, the importance ofthis new taxon warrants description and formal naming.

Description. The skull is rather large for its geological age,comparable to that of European Pachytragus (measure-ments: Table 1). Its most obvious feature is the strongcranial flexure (Fig. 2A). The angle between the fronto-parietal profile, behind the horn-cores, and that of the(missing) dorsal profile of the face, can be estimated at

about 100°; the angle between the basioccipital and thealveolar plane is about 50°. Correlated with this highlyderived skull flexure, the orbit is located far behind thetooth row, and the frontals are strongly elevated betweenthe horn-cores. The anterior part of the frontals, includingthe supra-orbital foramina, the nasals, the premaxillae,and most of the maxillae, are missing. There is a doublepostcornual fossa, which is in fact infracornual. All bonesof the low, broad braincase are fused, and no suture isvisible, but the parietal was certainly short; its dorsalprofile is slightly concave. The angle between it and theplane of the occipital is about 130°. The occipital has arounded outline, and is low and broad, with a centralsagittal crest (Fig. 2G). The mastoid and auditory areas are

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Figure 1. Map of Morocco with the location of Skoura in the Ouarzazate basin. Elevation data from http://hydrosheds.cr.usgs.gov/

Table 1. Measurements (in mm) of holotype skull of Skouraia helicoides.

Length from occipital condyle to P2 (estimated) 193Length from occipital condyle to M3 118Minimum width of braincase 69Width of occipital 82+Height of occipital (from top of foramen) 39Width of basioccipital 28.6Bicondylar width 56Width over M3s 75Width over horn core pedicles 91Antero-posterior diameter of horn-core 47.4Transverse diameter of horn core 41.1Length of right horn core 260+Length M1-M3 55.8Length P3-P4 21

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indistinct. On the basioccipital, the usual pairs of anteriorand posterior tuberosities are hardly distinct; they are infact little more than elevations above a pair of rectangularplateaus, only slightly narrower anteriorly than posteriorly,and separated by a long narrow groove in the midline(Fig. 2F); there are no accessory stop facets for the atlas.The choanae reach the level of the posterior border of M3,and the lateral indentations almost reach the same level(Fig. 2F).

The horn-cores are inserted above the orbits, and are

strongly inclined backwards, as their basal part is parallelto the tooth row in lateral view, and their tips are moreventral than their base (Fig. 2A). Basal divergence is about90°, and slightly increases towards the tips. They arestrongly spiralled, but the axis of revolution of the horncore probably remains within the bone, thus forming aclosed spiral. Torsion is homonymous, i.e. the right hornhas a clockwise torsion (its is perhaps necessary to stresshere that, although imperfectly preserved, the skull iswholly devoid of reconstruction, and therefore no error

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Figure 2. Skouraia helicoides sp. nov., holotype cranium. A, right lateral view; B, dorsal view; C, central view; D, posterior view; E, left tooth row P3–M3;F, stereo view of the cranial base; G, occipital view. Scale = 5 cm for Fig. E, 10 cm for Figs F, G, 20 cm for Figs A–D.

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can have occurred in fitting the horn-cores onto the skull).They have a strong anterolateral keel, which remainsstrong all the way to the tip, and a weaker anteromedialone; the anterior surface is almost flat between these keels,and the remaining part of the surface is rounded, withonly a slight mediolateral compression. Owing to imper-fect preservation of these parts, we do not know whetherthe base of the horn-cores and/or the frontal, werepneumatized.

The premolars are very short compared to the molars(Fig. 2E). The P4 has a rather square outline, being almostas long lingually as labially, and has a moderate, centrallyplaced, labial rib. This rib is much more mesially placed,and stronger, on P3, which is distinctly smaller than P4,making unlikely that the P2, now missing, was larger.Assuming that it was as large as P3, the index L P2-P4/LM1-M3 100 can be estimated at about 55, a very low value,comparable to that of modern Capra. The molars have noentostyles. The paracone has a weak labial rib, but there isnone on the metacone, whose labial wall is a flat depressedsurface between prominent styles; the metastyle of M3 isremarkably salient. On the whole, the dentition is there-fore clearly of the aegodont type.

Comparisons. The most remarkable feature of this newbovid is the homonymous direction of horn torsion. Mostspiral-horned bovids, living and fossil, instead have hornswith heteronymous torsion, i.e. the right horn-core has ananticlockwise torsion. This is true, in particular, of thoseantelopes with the most conspicuously spiralled horns,the African Tragelaphini, but also of the modern Antilope(Antilopini) and Addax (Hippotragini), whose horns aremarkedly spiralled, and of Kobus (Reduncini), Damaliscus(Alcelaphini) and Aepyceros, where torsion is muchweaker. Most of the abundant spiral-horned forms of thePalearctic late Miocene, such as Palaeoreas, Prostrepsiceros,Protragelaphus, and related genera, also have heteronymoustorsion. Thus, their few representatives in Africa, such asProstrepsiceros libycus Lehman & Thomas, 1987 fromSahabi (referred to Dytikodorcas by Bouvrain & Bonis,2007), or the ‘Palaeoreas’ from the Samburu Hills (Nakayaet al. 1987), can be readily excluded from the comparison.

A slight homonymous torsion can be found in manyAlcelaphini, especially Connochaetes, Megalotragus andNumidocapra, but no member of this tribe has horns thatare strongly spiralled, and they have a characteristic toothmorphology quite distinct from that of Skouraia (althoughthey share the same premolar reduction). It is also detect-able, but much weaker than in Skouraia, in some otherAfrican bovids that are clearly unconnected to genus,such as Menelikia and Antidorcas.

Bovids with a clear homonymous horn core torsion canbe divided into four groups for convenience, and comparedin the same order:1) the Oioceros group, mostly of late Miocene age;2) the Sinotragus group, also of late Miocene age, perhaps

related to the next group, or to the Pachytragus-Protoryxgroup;

3) many modern Caprini, including the sheep and somegoats;

4) a few fossils from northwestern Africa.

1) Among the informal Oioceros group, Hispanodorcas fromSpain (Thomas et al. 1982; Alcalá & Morales 2006) andGreece (Bouvrain & Bonis 1988) has slender, almoststraight horn-cores with weak torsion and no keels, whichare uprightly inserted far from the occipital, and arounded braincase. Hispanodorcas is clearly unrelated toSkouraia.

Pontoceros Verescagin et al., 1971, from the Plio-Pleistoceneof the Black Sea and Greece (Verescagin et al. 1971;Kostopoulos 1997) also has almost straight horn-cores, notspiralled like those of Skouraia, with a more triangularcross-section underlined by distinct furrows, and they aremuch less divergent.

Systematics of the late Miocene Oioceros-Samotragus-Samodorcas group is still debated, and its discussion isbeyond the scope of this paper. ‘Oioceros wegneri’ differsfrom Oioceros sensu stricto, and approaches Skouraia, in itsstronger cranial flexure and larger size, but differs fromSkouraia in several cranial features: the frontal is not stronglyelevated between the horn-cores; even the largest ‘O. weg-neri’ remain of moderate size; the basioccipital has verystrong anterior and posterior tuberosities; the occipitalcondyles are less broad relative to the occipital width. Thehorn-cores also offer clear distinguishing features betweenthe Oioceros group and Skouraia. In the Oioceros group, thesurface of the horn-cores is not smooth; instead, it isalways marked by one or more longitudinal groove(s) andadjacent or intervening ridge(s) that may be acute andbecome keel-like, but these keels result from the depres-sion of the areas adjacent to them, rather than from theirown raising above the horn core surface, so that thecross-section is quite different; in Oioceros sensu stricto, it isoften comma-shaped. Oioceros is also clearly unrelated toSkouraia.

2) Prosinotragus and Sinotragus from the Upper Miocene ofChina (Bohlin 1935), Samos (Solounias 1981), and Turkey(Geraads et al. 2002) have horn-cores that are short, onlyslightly divergent and close to each other at the base, withat most an incipient torsion, and with a keel that isanteromedial rather than anterolateral when present (inthe Samos and Turkish forms). All these features differfrom those of Skouraia helicoides.

The Pachytragus-Protoryx group has been revised byKostopoulos (2009), who considered both names as syn-onymous and created the new genus Skoufotragus. Thisgroup, best known from the Turolian (Gentry 2003, andreferences therein), had long been placed among theHippotragini, until Gentry (1971, 2000) convincinglyargued that it is closer to the Caprini. Kostopoulos(2005, 2009) maintained that its affinities are still open todiscussion because the i1 is larger than i2, unlike Caprini,but the size of i1 seems to be unrelated to phylogeny, andwe suspect that its small size in modern Caprini is a conse-quence of its incipient hypsodonty, so that a moderatelylarge i1 might just be primitive. Members of the Pachy-tragus-Protoryx group have a derived skull with a longface, a posteriorly located orbit, a strong cranial flexure,rather simple horn-cores variably curved backwards butwithout torsion, and an oval cross-section without keels.

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Identification of the dentitions at species level is oftendifficult, but many of them resemble Skouraia in theiraegodont pattern, with short premolars, and molars withweak labial ribs and flat metacone, prominent styles, noentostyles, and often a central fossette. Pachytragussolignaci Robinson, 1972, from the middle/late Miocene ofTunisia is the only African form; it resembles Skouraia inthe slight spiralling of a horn-core figured by Robinson(1972, Fig. 3) but otherwise clearly differs in the strongtransverse compression of the horn-cores. Horn-coremorphology rules out any close connection betweenSkouraia and this group.

3) In many modern Caprini, such as Capricornis,Naemorhedus, Oreamnos, Pseudois, and Rupicapra, the horn-cores are short, conical, and lack keels and torsion, so thatthey differ completely from those of Skouraia; those ofHemitragus have keels but are also short and simple; thoseof the ‘Ovibovini’ Ovibos and Budorcas are very peculiarand completely different, although a clear homonymoustorsion can be found in Euceratherium, an extinct relativeof the musk-ox. Ammotragus, Capra and Ovis deservegreater attention. They have highly derived skulls with along face and an orbit located posteriorly, and a stronglyflexed cranium. Their horn-cores display a wide range ofshapes; they are always curved, usually strongly so, andoften show some spiralling. In Capra and Ammotragus, it isonly incipient (except in C. falconeri, whose horn-cores aretightly twisted), and usually heteronymous, but some(especially domestic forms) may have a weak homonymoustorsion. Many Capra (especially C. falconeri and domesticforms), as well as the probably related Bouria from theEthiopian Pleistocene (Vrba 1997), have a strongly com-pressed cross-section with an anterior keel; Caprawodaramoya Bibi et al. 2012, from the early Pleistocene ofEthiopia is similar but without a clear anterior keel. OtherCapra, and Ammotragus, have a more triangular cross-section, with rounded angles, and a tendency to form aflattened anterior face, limited by a poorly indicatedanteromedial change of curvature, which may become akeel near the tip, and a better indicated anterolateral one,which may almost be a keel (e.g. in C. sibirica). In Ovis, thecross-section is similar but often less compressed andwithout true keels; however, the greater basal divergencethat increases above the base and the strong homonymoustorsion are resemblances with Skouraia. Thus, amongliving bovids, it is undoubtedly with the Caprini that wefind any possible similarities with Skouraia.

4) In Northwestern Africa, Parantidorcas latifronsArambourg, 1979 from the Pliocene of Aïn Brimba inTunisia has horn-cores that have a weak but distinctheteronymous torsion, but this is a much smaller speciesthat probably belongs to the Antilopini.

The northwestern African fossil form most similar toSkouraia is Benicerus theobaldi Heintz, 1973, from BeniMellal in Morocco. This site has traditionally been assignedto the middle Miocene because it lacks hipparions, andJaeger (1977) even concluded that it is not immediatelyanterior to the Vallesian because another site, Pataniak 6,is also ante-Vallesian but younger than Beni Mellal.

However, Ginsburg (1977), followed by Werdelin &Peigné (2010), noted that its carnivore fauna is similar incomposition to late Miocene ones; we estimate the age ofBeni Mellal at c. 12 Ma.

The holotype, and only described specimen, of B. theo-baldi is an incomplete left horn-core with most of the orbit(Heintz 1973, pl. 1); unfortunately, we could not locate it inthe Muséum National d’Histoire Naturelle, Paris. FromHeintz’ description and figures, this horn-core is stronglycurved posteriorly, moderately compressed transverselybut with a compression that quickly increases above thebase; it has a clear anticlockwise torsion, and a sharpanterior keel. It differs from the type of S. helicoides in itssmaller size, shortness, more upright insertion, greaterapical compression, and complete lack of an anteromedialkeel and of an anterior surface. Benicerus is too poorlyknown for in-depth comparisons with Skouraia, but theyare the only Miocene African bovids with distinctlyhomonymous torsion, and both are from Morocco.

There are two or three bovid species in the Upper Mio-cene of Jebel Krechem in Tunisia (Geraads 1989); somehorn-cores could belong to a form close to Prostrepsiceros,but there are also some fragments of spiralled horn-coreswhose direction of spiralling cannot be determinedbecause they lack the base, but one of them looks sosimilar to the type of B. theobaldi that it is hard to believethat torsion was not homonymous, as in this species. Thecross-section is ovoid, transversely compressed, with akeel that must have been anterior, and a hint of aposteromedial one. The presence of this second keel is theonly difference with B. theobaldi, and we consider likelythat a species close to that of Beni Mellal was also presentat Jebel Krechem, whose age is probably equivalent toEuropean MN10. Unfortunately, no spiral-horned bovidhas been found in the Beglia Formation of Tunisia (Robin-son 1986), which is of intermediate age, but it is conceiv-able that Benicerus survived into the late Miocene. Howclosely this genus is related to Skouraia is hard to tell; aBenicerus theobaldi–Skouraia helicoides lineage can behypothesized, but unsubstantiated on the basis of thevery poor record from Beni Mellal, and we prefer topropose Skouraia as a distinct genus.

Behaviour. The strong divergence of the horn-cores, andtheir strong inclination, with no part of them being dorsalto the frontal plane, strongly suggest that Skouraia per-formed some kind of the agonistic fighting behaviourknown as ramming or Rammkampf, in which the oppo-nents violently clash their heads after charging. It is bestknown in the Caprini Ovis and Ammotragus, in Ovibos, butalso occurs in Connochaetes and some Bovini. SinceSkouraia is obviously unrelated to any of the latter taxa,this inferred behaviour may be taken as supporting anassignment of this new taxon to the Caprini.

CONCLUSIONThe most likely scenario is that Skouraia is an early,

Ovis-like, offshoot of the Caprini, possibly related toBenicerus. Phylogenetic relationships between the livingmembers of this tribe have recently been investigated,with limited consistency in the results (Schafer & Hall

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2010; Bibi et al. 2012, and referencess therein). Moleculardating suggests that the early diversification within thetribe started in the late Miocene (Ropiquet & Hassanin2005). The fact that Myotragus, which does not occupy abasal position in this group (Lalueza-Fox et al. 2005), musthave reached the Balearics during the Messinian, impliesthat this is a minimum date, and if the shortenedmetapodials of Aragoral mudejar, of MN10 age, are indeeda synapomorphy with the Caprini (Alcalá & Morales1997), the divergence of this tribe from the Hippotraginiand Alcelaphini might go back to the middle Miocene(Bibi et al. 2009). It is therefore entirely possible that theNorth African latest Miocene witnessed an early diversifi-cation of the Caprini, which filled there a niche occupiedin Eurasia by a variety of other antelopes. It is also likelythat this group settled earlier and more deeply in Africathan hypothesized by Bibi et al. (2012), who regard itsspotty record in this continent as mere successive wavesof migrations from Eurasia.

We are especially grateful to B. Tahiri for having allowed us to study the specimenin his private museum. Thanks to A.W. Gentry and to an anonymous reviewer fortheir useful comments that significantly improved the manuscript. Thanks also forgiving access to collections in their care to J. Lesur-GebreMariam and C. Argot(Muséum National d’Histoire Naturelle, Paris), and M. Bertling (Geologisch-Paläontologisches Institut, Universität Münster).

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HEINTZ, E. 1973. Un nouveau bovidé du Miocène de Beni Mellal,Maroc: Benicerus theobaldi n.g., n.sp. (Bovidae, Artiodactyla,Mammalia). Annales Scientifiques de l’Université de Besançon 18, 245–248.

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KOSTOPOULOS, D.S. 1997. The Plio-Pleistocene artiodactyls(Vertebrata, Mammalia) of Macedonia. 1. The fossiliferous site‘Apollonia-1’, Mygdonia basin of Greece. Geodiversitas 19, 845–875.

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Morphometric analysis of modern human crania: aframework for assessing early Pleistocene hominids

E.J. Odes1,2 & J.F. Thackeray1*1Institute for Human Evolution, University of the Witwatersrand, P.O. WITS, Johannesburg, 2050 South Africa

2School of Geosciences, University of the Witwatersrand, P.O. WITS, Johannesburg, 2050 South Africa

Received 16 January 2012. Accepted12 October 2012

INTRODUCTIONThackeray (2005) has previously examined cranial varia-

tion in modern hominoid primates, excluding modernhumans, in the context of variation in skulls of extinctAfrican hominins. In this study we obtain cranial datafrom a sample of modern humans, using more than100 landmarks, to provide a frame of reference for assess-ing Plio-Pleistocene hominins.

MATERIALS AND METHODSThe modern comparative human sample in the Dart

Collection at the University of the Witwatersrand, fromcadavers, includes 24 African crania: Ndebele (two male,two female), Shangaan (two male, two female), Sotho (twomale, two female), Swazi (two male, two female), Tswana(one male, one female), Xhosa (two female, two male),Zulu (one male), and one African human cranium; andfive crania catalogued as Europeans (Caucasian). Thechoice of sample size was arbitrary, with the objective ofsecuring a total sample of at least 25 individuals, recogniz-ing that this represents only part of the range of variationthat would be expected from a global sample of H. sapiens.

The measurements on the modern crania (Table 1) werebased on landmarks previously used in Wood’s (1991)study of fossil hominins. The landmarks includeProsthion (pr); Nasospinale (ns); Nasion (n); Glabella (g);Bregma (b); Vertex (v); Apex (ap); Lambda (l);Opistocranion (op); Inion (i) Opisthion (o); Basion (ba);Alveolon ( alv); Staphylion (sta); Orale (ol); Euryon (eu);Porion (po); Mastoidale (ms); Alare (al); Orbitale (or);Zygion (zy); Ectoconchion (ec); Ectomalare (ecm); Pterion(pt); Endomalare (enm); Frontotemporale (ft).

Measurements were obtained using (1) digital callipersmanufactured by the Mitutoyo Corporation (product

name Digimatic Caliper; model no: CD-6 inch CX; codeno: 500-171-20; serial no: 09093312; measuring range0–150 mm; minimum indication 0.01 mm); (2) a two-button digital calliper manufactured by Mitutoyo Corpo-ration, with the same technical specifications (resolution:0.01 mm); and (3) a Mitutoyo digital linear spreadingcalliper with a 300 mm digital scale and a throat depth of150 mm. The modern crania were positioned on a foambase for stability and protection. Measurements weretaken using the same calipers throughout the study tominimize measurement error. Curved regions weremeasured using chord distance.

Statistical methodThe method that is used in this study has been devel-

oped by Thackeray et al. (1997, 2007). Morphological varia-tion within a species can be quantified using least squareslinear regression analysis of measurements of pairs ofspecimens. The degree of similarity between two speci-mens of the same species can be expressed by comparingmeasurements of a reference specimen A (associated withthe x-axis) and conspecific specimen B (associated withthe y-axis). In such cases there is generally little scatteraround the regression line, associated with the linearregression equation y = mx + c, where m represents theslope of the regression line, and c represents the constant;the log transformed standard error of the slope m isreferred to as log s.e.m. The limited scatter around theregression line is associated with similarity in shape of thetwo conspecific specimens.

The degree of scatter around the regression line isquantified by the standard error of the m-coefficient (s.e.m).

The s.e.m value for pairs of conspecific pairs is relativelylow. By contrast, when measurements of two specimensrepresenting two different species are compared, there is

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*Author for correspondence. E-mail: [email protected]

Craniofacial measurements have been obtained from modern human skulls from cadavers representing several southern Africanpopulation groups including Ndebele, Shangaan, Sotho, Swazi, Tswana, Xhosa and Zulu, in addition to European Homo sapiens. Themeasurements were obtained from crania in the Dart Collection housed at the School of Anatomical Sciences of the University of theWitwatersrand. Pairwise comparisons, using least squares linear regression analysis of cranial measurements, were used to calculate thestandard error of the m-coefficient associated with the general equation y = mx + c, where m is the slope of the regression line. The stan-dard error of the m-coefficient is a measure of the degree of similarity between specimens. Log transformed s.e.m values (log s.e.m) showa normal distribution with a mean value of –1.84 ± 0.087 (n = 384 pairwise comparisons). These results can be used as a frame ofreference for comparing Early Pleistocene specimens. For example, a comparison between KNM-ER 1813 (attributed to H. habilis) andKNM-ER 3733 (attributed to H. erectus or H. ergaster) is associated with a log s.e.m value of –1.844. Despite differences in size, these twopenecontemporary hominid fossils are associated with a high probability of conspecificity, since the log s.e.m value is identical to themean log s.e.m value of –1.84 obtained for pairwise comparisons of modern Homo sapiens.

Keywords: Homo sapiens, Homo ergaster, Homo habilis, morphology, craniofacial.

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a greater degree of scatter around the regression line andthe s.e.m value is relatively high (Thackeray et al. 1997;Aiello et.al., 2000). The distribution of s.e.m values obtainedfrom pairwise comparisons of extant conspecific pairs ofspecimens has been examined. Thackeray et.al. (1997)showed there is a log normal distribution of s.e.m valueswhen pairwise comparisons are made between conspecificspecimens of extant vertebrates including mammals,birds, reptiles and extant invertebrates. In a larger study(Thackeray, 2007), log s.e.m values are inclined to becentrally distributed around a mean log s.e.m value of–1.61 ± 0.23 (including vertebrates and invertebrateswhere n = 1424 specimens), not significantly differentfrom a mean log s.e.m value of –1.66 ± 0.20 for inverte-brates (n = 172 specimens).

Thackeray (2007) proposed that when comparisons aremade between any two specimens of the same species,

the log s.e.m approximates a ‘biological species constant’(T = –1.61) which is considered to prevail over evolution-ary time and geographical space. The advantage of thisapproach is that the mean log s.e.m value of –1.61 ± 0.23facilitates a definition of a species based on morphometricanalysis. It allows for the assessment of probabilities ofconspecificity associated with fossil specimens includinghominins from South and East Africa, taking into accountthe fact that there is morphological variation in time andspace, and recognizing that there is no clear boundarybetween hominin species in space and time, and no clearboundary between Australopithecus and Homo. Themethod has been used in this study to assess the degree ofsimilarity between a sample of modern specimens ofHomo sapiens, representing only part of the range of varia-tion that would be expected in a global sample. As anexample of the application of this approach, comparisons

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Table 1. List of craniofacial measurements.

Number Measurement Number Measurement

1 Glabella–opisthocranion2 Posthion–inion3 Posterior cranial length4 Basion–bregma5 Basion–nasion6 Porion height7 Mastoid length8 Minimum frontal breadth9 Maximum parietal breadth10 Maximum temporal breadth11 Biporionic breadth12 Supramastoid breadth13 Maximum breadth across mastoid process14 Biaterionic breadth15 Interentoglenoid breadth16 Entoglenoid breadth17 Glabella–bregma18 Glabella–bregma19 Postglabellar sulcus–bregma20 Postglabellar sulcus–bregma21 Bregma–left pterion22 Bregma–left pterion23 Bregma–right pterion24 Bregma–right pterion25 Parietal sagittal length26 Parietal sagittal length27 Parietal temporal length28 Parietal temporal length29 Parietal coronal breadth30 Parietal coronal breadth31 Parietal lambdoid length32 Parietal lambdoid length33 Bregma–asterion34 Bregma–asterion35 Lambda–inion36 Lambda–inion37 Inion–opisthion38 Inion–opisthio39 Occipital sagittal length40 Occipital sagittal length41 Biasterionic breadth42 Biasterionic breadth43 Superior facial height44 Superior facial length45 Alveolar height46 Subnasale–prosthion47 Subnasale–prosthion (horizontal projection)48 Subnasale–prosthion (vertical projection)49 Superior facial breadth50 Biorbital breadth51 Bijugal breadth

52 Bizygomatic breadth53 Bimaxillary breadth54 Outer alveolar breadth55 Anterior interorbital breadth56 Orbital breadth57 Orbital height58 Orbitale–zygomaxillare59 Minimum malar height60 Malar thickness61 Width temporal gutter62 Vertical thickness of supraorbital torus63 Anteroposterior thickness of supraorbital torus64 Anteroposterior thickness of supraorbital torus65 Anteropoterior thickness of glabella66 Frontal torus breadth67 Frontal torus breadth68 Maximum nasal width69 Nasal height70 Rhinion–nasospinale71 Sagittal length of nasal bones72 Superior breadth of nasal bones74 Inferior breadth of nasal bones75 Infratemporal fossa depth76 Foramen magnum length77 Foramen magnum maximum width78 Occipital condyle maximum length79 Occipital condyle maximum width80 Mandibular fossa length81 Mandibular fossa length82 Mandibular fossa breadth83 Mandibular fossa breadth84 Mandibular fossa depth85 Depth of postglenoid process86 Depth of articular eminence87 Maxillo-alveolar length88 Maxillo-alveolar breadth89 Palate length (orale-staphylion)90 Palate length (orale-palatomaxillary suture)91 Palate breadth92 Incisive canal-palatomaxillary suture93 Internal alveolar breadth at M394 I1-I2 alveolar length95 Canine alveolus breadth96 P3-P4 alveolar length97 M1-M3 alveolar length98 Intercanine distance99 P3 interalveolar distance100 P4 interalveolar distance101 M2 interalveolar distance102 M3 interalveolar distance103 Palatal height

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are between two penecontemporary Early Pleistocenehominin specimens, KNM-ER 1813 (attributed to H.habilis) and KNM-3733 (attributed to H. ergaster or H. erectus),dated around 1.6 million years, from Koobi Fora (EastTurkana) in Kenya (Wood, 1991).

RESULTSResults obtained from pair-wise comparisons of crania

of modern humans (Table 2) are presented in Fig. 1.

COMPARISON BETWEEN FOSSIL CRANIAA log s.e.m value of –1.84 has been obtained from a

comparison of two early Pleistocene hominid crania,KNM-ER 1813 (attributed to H. habilis) and KNM-ER 3733(attributed to H. ergaster or H. erectus).

DISCUSSION AND CONCLUSIONThe results of this study show that the log s.e.m values

obtained from 384 pairwise comparisons of human craniadisplay a normal distribution, similar to results obtainedfrom previous studies of both vertebrates and inverte-brates (Thackeray et al. 1997; Thackeray 2007). The meanlog s.e.m value of –1.84 ± 0.09 obtained from the humancrania provides a frame of reference for assessing EarlyPleistocene hominids.

Thackeray et al. (1997) compared conspecific specimensof extant vertebrates including mammals such as primates,rodents and ungulates; birds; reptiles; and invertebrates(Coleoptera and Lepidoptera). In a preliminary studythe mean log-transformed s.e.m value calculated byThackeray et al (1997) from comparisons of measurementsof conspecific pairs of vertebrates and invertebrates was–1.78 ± 0.27 for 1260 specimens (70 extant species).Extending this approach Thackeray (2007) provided astatistical definition of a species, taking advantage of asubstantially larger sample of vertebrates. Using thatenlarged sample Thackeray (2007) calculated a meanlog s.e.m of –1.61 ± 0.23 (n = 1424 specimens), which isnot significantly different from the mean value obtainedfrom the initial study.

The result of –1.84 obtained from Homo sapiens crania inthis study is lower than the mean log s.e.m value of –1.78(Thackeray et al. 1997) and –1.61 (Thackeray, 2007)obtained for large samples of a diversity of vertebratesand invertebrates.

Despite differences in size, KNM-ER 1813 and KNM-ER3733 are examples of early Pleistocene hominid fossilswhich are associated with a high degree of similarity. Thelog s.e.m. value of –1.84 obtained from the comparison ofthese two specimens is identical to the mean log s.e.m.

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Figure 1. Normal distribution of log s.e.m values obtained from pair-wise comparisons of Homo sapiens crania in the Dart Collection. The meanlog-transformed s.e.m value based on comparisons of conspecific pairs of Homo sapiens is –1.84 ± 0.09 ( n= 384 comparisons).

Table 2. Human specimens from the Dart Collection, School of Anatomi-cal Sciences, University of the Witwatersrand. This list includes cata-logue numbers, population affinities, age and sex.

Catalogue Population Age Sexnumber affinity

4011 Tswana 54 m4069 Tswana 20 f3955 Zulu 56 m1532 Ndebele 69 f1549 Ndebele 28 f1274 Ndebele 39 m1535 Ndebele 35 m172 Shangaan 60 m58 Shangaan 38 m263 Shangaan 30 f3057 Shangaan 30 f2492 Sotho 21 f2307 Sotho 60 f2248 Sotho 48 m2077 Sotho 48 m1360 Swazi 19 f1534 Swazi 24 f2014 Swazi 42 m1362 Swazi 49 m1333 Xhosa 48 m400 Xhosa 36 m22 Xhosa 30 f761 Xhosa 37 f4035 African 22 f2179 Caucasian 19 f3902 Caucasian 40 m3129 Caucasian 29 f2186 Caucasian 57 m3545 Caucasian 40 m

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value of –1.84 obtained for pairwise comparisons ofmodern Homo sapiens. The implication is that there is ahigh probability that KNM-ER 1813 and KNM-ER 3733 areconspecifics, despite the fact that they have previouslybeen attributed to different species of Homo (H. habilisand H. erectus/ergaster). This serves as one exampledemonstrating the applicability of a morphometricapproach to assess probabilities of conspecificity in fossilhominids.

Thackeray (2005) has previously suggested thatKNM-ER 1813 is a small female of a species also repre-sented by KNM-ER-3733, considered to be a large male. Bothare dated to about 1.6 million years before present (BP).

We are grateful to the NRF, the University of the Witwatersrand, the AndrewMellon Foundation, and the French Embassy in South Africa for financial support.

We thank Brendon Billings for access to material in the Dart Collection. We alsothank Alan Morris and an anonymous referee for helpful comments.

REFERENCESTHACKERAY, J.F. 2005. Probabilities of conspecificity: an application of a

morphometric approach to extant and extinct hominoids. In: �trkalj,G., Pather, N. & Kramer, B. (eds), Voyages in Science. Essays by SouthAfrican Anatomists in Honour of Phillip V. Tobias’s 80th Birthday, 85–97.Pretoria, Content Solutions.

THACKERAY, J.F., 2007. Approximation of a biological species constant?South African Journal of Science 103, 489.

THACKERAY, J.F., BELLAMY, C.L., BELLARS, D., BRONNER, G.,CHIMIMBA, C., FOURIE, H., KEMP, A., KRUGER M., PLUG, I.,PRINSLOO, S., TOMS, R., VAN ZYL, A.J., & WHITING, M.J. 1997.Probabilities of conspecificity: application of a morphometrictechnique to modern taxa and fossil specimens attributed toAustralopithecus and Homo. South African Journal of Science 93 : 195–196.

WOOD, B. 1991. Koobi Fora Research Project, Vol. 4. Hominid Cranialremains. Oxford, Clarendon Press.

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Palaeontological Society of Southern Africa16th Biennial Conference, University of Cape Town

5–8 September 2012(An index to authors is given on page 58)

Basal therocephalian from the Middle Permianof South Africa: taxonomy and geographic andtemporal distribution

Abdala, Fernando; Day, Michael; Rubidge, BruceBernard Price Institute for Palaeontological Research, Palaeosciences Centre,

University of the Witwatersrand, East Campus, Private Bag 3, WITS 2050,Johannesburg, South [email protected]

Important changes in amniote taxonomic representationoccurred during the Middle Permian, the most significantbeing an explosive emergence of therapsids. The MiddlePermian tetrapod record of the South African KarooSupergroup has the oldest advanced theriodonts, includ-ing gorgonopsians and therocephalians. The oldesttherocephalians are the largest members of this lineageand clearly were top predators in the Middle Permian.Two main basal therocephalian families are present at thisage and can readily be distinguished by snout propor-tions: Scylacosauridae with a long and comparativelynarrow snout, and Lycosuchidae with a short and broadsnout. New findings of representatives of these twogroups of basal therocephalians in the Karoo Basinprompted a revision of the taxonomy, and geographic andtemporal distribution in the Karoo Basin. In theEodicynodon Assemblage Zone (AZ), the lowermostbiozone of the Beaufort Group, they are represented bytwo scylacosaurid taxa. The overlying Tapinocephalus AZ,by far the thickest vertebrate biozone of the Group, showsa great increase in diversity and abundance of basaltherocephalians. Thus from only two specimens repre-sented in the Eodicynodon AZ, more than 100 specimensare known from the Tapinocephalus AZ. The last record ofbasal therocephalians is represented in the PristerognathusAZ in which the diversity and abundance is clearly dimin-ished. Lycosuchids are generally considered in phylogeneticanalyses as the most basal therocephalians; however, theirfossil records (apart from three uncertain localities) are re-stricted to the upper Tapinocephalus and the overlyingPristerognathus Assemblage Zones. The most advancedscylacosaurids are known from the basal Eodicynodon AZuntil the Pristerognathus AZ. This revision will allow explo-ration of the actual extension of the first important diversi-fication of therocephalians, and will provide a betterknowledge of the distribution, temporal and geographic,of these early representatives of therocephalians in theKaroo Basin.

Ontogeny of the Early Triassic Thrinaxodonliorhinus (Therapsida, Cynodontia): dentalmorphology and replacement

Abdala, Fernando1; Jasinoski, Sandra2; Fernandez, Vincent1

1 Bernard Price Institute for Palaeontological Research, Palaeosciences Centre,University of the Witwatersrand, East Campus, Private Bag 3, WITS 2050,Johannesburg, South [email protected]

2 Vancouver, British Columbia, Canada. Former postdoctoral fellow in the ZoologyDepartment, University of Cape Town, Rondebosch, South [email protected]

We present a detailed study on variation of the dentalmorphology and replacement in the Early Triassiccynodont Thrinaxodon liorhinus. For this study we analysedfive specimens ranging from 37–87 mm in skull lengthusing micro computed tomography (microCT) scanningtechniques, which were supplemented by detailed ana-tomical analysis of 41 specimens with a basal skull lengthof 30–96 mm. Our results confirm the alternate replace-ment of the postcanines and the posterior migration of thepostcanine series (including the loss without replacementof the anteriormost postcanines). Even when most of theobservations point to a posterior-to-anterior replacementwave, the evidence is not clear-cut. Lower postcanines aremore numerous and present a more complex morphologythan the upper postcanines, even in the same individual;only the lower postcanines have more than three sectorialcusps and a cingular collar on the lingual margin.Complexity of the postcanines increases from the smallestindividual to specimens of 75 mm of skull length, butcomplexity decreases in larger specimens. The virtualextraction of functional and replacement teeth permittedus to conclude that in most of the cases, the upper canineswere replaced anteriorly while lower canines were replacedposteriorly. The presence of two simultaneous replacementsof the upper canine tooth was observed in two smalljuveniles, suggesting a higher rate of canine replacementat younger age. Incisors also had a sequential replacementpattern, and more replacement teeth were present inmedium-sized individuals.

New vertebrates from the Permian Pedra de FogoFormation, Parnaíba Basin, northeastern Brazil

Angielczyk, Kenneth1; Cisneros, Juan C.2;Mariscano, Claudia A.3; Richter, Martha4;Fröbisch, Jörg5; Kammerer, Christian F.5;Smith, Roger M.H.6; da Conceição, Domingas, M.2;de Castro Silva, Mayana2

1 Field Museum of Natural History, Chicago, [email protected]

2 Universidade Federal do Piauí, Piauí, [email protected] / [email protected]

3 CONICET Universidad de Buenos Aires, Buenos Aires, [email protected] / [email protected]

4 Natural History Museum, London, United [email protected]

5 Museum für Naturkunde, Humboldt-Universität zu Berlin, Berlin, [email protected] / [email protected]

6 Iziko South African Museum, Cape Town, South [email protected]

The Permian assemblage of the Pedra de Fogo Formation(PFF) includes petrified wood, fish, and a temnospondylamphibian. The assemblage has received little attention,however, and its precise age is uncertain, with estimatesspanning most of the Permian. In 2011 and 2012 we con-

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ducted fieldwork in the Parnaíba Basin (Maranh o, Piauíand Tocantins states), to better understand the chronolog-ical and palaeoenvironmental contexts of the PFF fauna.

The PFF accumulated in a large shallow intra-continentaldepression, the Parnaíba Basin. Palaeoenvironmentschanged with progressive aridification and continentality,from a large shallow epeiric sea that supported a flourish-ing chondrichthyan community, to isolated continentalwater bodies between aeolian dune fields.

Fish are the most abundant vertebrate fossils. Chon-drichthyes dominate in wavy-bedded sandstones andsiltstones that accumulated towards the basin’s depo-centre. Ctenacanths, xenacanths, and other chondrich-thyans include new taxa. The holocephalan (petalodont)Itapyrodus puctatus proved ubiquitous in the nearshoreand evaporitic shoreline facies, where teeth of Aniso-pleurodontis pricei, a large, endemic, shark-like predatorare also common. In Tocantins, we collected specimensof the shark Glikmanius, also known from the Lower Perm-ian of the U.S.A. and the Carboniferous of Russia andEurope. Dipnoan fossils are relatively rare, but we tenta-tively recognize at least four distinct tooth platemorphologies. In more continental deposits in the upperPFF, we collected osteichthyans (lower actinopterygiansand coelacanthiforms) with close affinities to Mesozoictaxa.

The putative archegosaurid Prionosuchus is the onlytetrapod previously known from the PFF. We have addi-tional amphibians from two areas, representing at leastthree new taxa. Specimens from the eastern edge of thebasin show some affinities with tupilakosaurids, whereasthose from the more central part of the basin may repre-sent brachyopids and rhitidosteids, all best known fromthe Triassic. During the Permian the Parnaíba Basin waslocated in the equatorial regions of Pangaea, comparableto the Moradi Formation of Niger, which includes ananachronistic mixture of taxa that may have been commonin central Pangaea. The PFF appears to include a mix ofclades best known from the Early Permian and Mesozoic.The PFF has the potential to yield new insights intoPermian biogeography and the end-Permian extinctionand recovery.

Improved biostratigraphic correlations betweenUpper Permian rocks in the Karoo, Luangwa andRuhuhu Basins: implications for biogeography andthe end-Permian extinction

Angielczyk, Kenneth1; Sidor, Christian2;Steyer, Jean-Sebastien3; Smith, Roger4

1 Department of Geology, Field Museum of Natural History, Chicago, [email protected]

2 Burke Museum and Department of Biology, University of Washington, Seattle,U.S.A.

3 Department Earth History, CNRS-Museum national d’Histoire naturelle Paris,France

4 Department of Karoo Palaeontology, Iziko South African Museum, Cape Town,South Africa

Upper Permian tetrapod fossils were discovered in theKaroo Basin in the mid-19th century, but similar faunalassemblages were only found in the Luangwa (Zambia)and the Ruhuhu (Tanzania) Basins in the early 20th century.The number of faunal assemblages present in the latter

basins and their correlation with the Karoo are uncertain,with two to three assemblages typically recognized in theUsili (formerly Kawinga) Formation of Tanzania and up tofour in the Upper Madumabisa Mudstone of Zambia. Avariety of correlations with the Karoo have been proposed,with both basins potentially preserving time equivalentsof the Pristerognathus to Dicynodon Assemblage Zones.Taxonomic confusion surrounding key groups, such asdicynodont therapsids, has hampered the developmentof more refined correlations.

Since 2007 we have worked in the Luangwa andRuhuhu Basins. These data, combined with those fromearlier collections and recent taxonomic revisions, pro-vide new insights into these Upper Permian faunas.Co-occurrence of species indicates that the Usili Forma-tion and the Upper Madumabisa Mudstone each host asingle Permian assemblage, although their preservationvaries; e.g. specimens from the basal conglomerate unit ofthe Usili Formation tend to be fragmentary and encrustedwith a thin hematite rind, whereas those higher in thesection are more complete. A Middle Permian-age fauna isalso present in the Ruhuhu Basin Ruhuhu Formation.

Stratigraphically long-ranging taxa like Dicynodontoidesor Pristerodon are present in the Upper MadumabisaMudstone and the Usili Formation, but the occurrence ofmore restricted taxa such as Odontocylops and Kitching-anomodon in the Luangwa Basin strongly suggests a corre-lation between the Cistecephalus Assemblage Zone of theKaroo and the Madumabisa assemblage. Dicynodon hueneiand Katumbia in both basins suggests that the Usili assem-blage also correlates with the Cistecephalus Zone.

Revised faunal lists for the Cistecephalus Zone, the UpperMadumabisa Mudstone, and the Usili Formation showthat each has similar proportions of endemic species. TheMadumabisa fauna, however, is more similar to thatCistecephalus Zone than to the Usili Formation, indicatingthat geographic proximity was not the sole factor control-ling distribution of these tetrapods. Ecological structureand taxonomic composition, with therapsids numericallyand taxonomically dominant, are shared. This diversitycontrasts strongly with the early Middle Triassic assem-blages from the same basins, suggesting that disruptionsassociated with the end-Permian extinction caused diver-gent recoveries.

Cretaceous and Tertiary gymnosperm andangiosperm woods from southern Africa

Bamford, MarionBPI Palaeontology, School of Geosciences, University of the Witwatersrand,

Private Bag 3, WITS 2050, South [email protected]

The Cretaceous and Tertiary fossil wood record forsouthern Africa is not extensive but there is a variety oftaxa from sites widely distributed over the southern partof the continent. Early Cretaceous woods are preservedonshore and offshore along the west coast, and onshorealong the east coast. These include woods of theAraucariaceae, Cheirolepidiaceae and Podocarpaceae, thelatter of which there are at least five species. Late Earlyand early Late Cretaceous woods from the east coast

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include a variety of Podocarpaceae woods as well as treeferns of Osmundacaulis species. This diversity of fernsextends into the Late Cretaceous and the gymnospermtaxa, although still of the Podocarpaceae, are mostlydifferent species from the older taxa. The most commonUpper Cretaceous species is Podocarpoxylon umzambenseSchultze-Motel and it was widespread in southern Africa.Some of these specimens have indistinct growth ringsand others have clear rings but variable in width. The cli-matic implications are explored further. From the UpperCretaceous Umzamba beds (southern coast) there arefossil logs of angiosperms: Monimiaceae and Euphorbia-ceae. Gymnosperm woods of the Araucariaceae andPodocarpaceae are also abundant from this locality.Tertiary woods are predominantly angiospermous. Thereis only one known Eocene locality and it has a mix ofwoods with well-defined family affiliations and someindeterminate woods. Oligocene woods are very rare butthere are abundant Miocene sites with wood and faunalremains. The common families in the fossil record are stillcommon today and include Anacardiaceae, Burseraceae,Combretaceae, Dipterocarpaceae, Fabaceae, Myrtaceaeand Oleaceae. These wood specimens do not have cleargrowth rings but their taxonomic affinities and woodstructure, such as size and density of vessel members, canbe used to infer local climate. There are no silicifiedgymnosperm woods from the Miocene but new recordsfrom the Cape region show the continued presence ofcarbonized Podocarpaceae. Today the only indigenousconifers in southern Africa are four species of Podocarpa-ceae and one of Cupressaceae.

A Cenozoic foraminiferal record and itsassociated palaeoenvironmental conditionsfrom the Namibian outer shelf

Bergh, Eugene1,2

1 Department of Geological Sciences, University of Cape Town2 Natural History Department, Iziko South African Museum, Cape Town.

[email protected]

The Benguela Current is one of four major easternboundary current regions worldwide where coastalupwelling dominates. The cold Benguela Current has amarked influence on the Namibian shelf and micro-faunal distributions. Depocentres may have shiftedbetween the inner and outer shelf over geologic time asattested by foraminiferal assemblages. Cenozoic-agedsediments from the Namibian phosphogenic beltbetween Lüderitz and Walvis Bay on the outer shelfrevealed three distinctive foraminiferal assemblagessuggesting three geological ages (Miocene, Pliocene andHolocene) and the absence of the Pleistocene record.Sixteen 1.8 m long cores, from a depth of 199 to 309 mexhibit a coarsening-upward sequence with an increase offoraminiferal abundance in the Holocene-aged units. Thelower units of the cores revealed the occurrence of theextinct families Stilostomellidae and Nodosariidaesuggesting an age of Miocene to Pliocene sedimentation.Uvigerina sp. together with Nodogenerina sp. andStilostomella sp. suggest Pliocene sedimentation. Com-plete mollusc shells Dosinia lupinus and Lucinoma capensis

confirm a Holocene age for the upper core units correlat-ing with the dominant foraminiferal species Ammoniajaponica and Elphidium advenum. The biostratigraphic agescan further be constrained with Sr-isotope data. Themicrofossil and mollusc occurrences indicate that bottom-water conditions had variable oxygen content throughoutgeologic time. Benthic foraminifera display an inversecorrelation with pelletal phosphorite components. Themicropalaeontology of the cores suggests a heteregeneousNamibian outer shelf with variable sedimentation anderosional rates.

Osteohistology of Triassic archosauromorphs inSouth Africa

Botha-Brink, Jennifer1; Smith, Roger2

1 Karoo Palaeontology, National Museum, P. O. Box 266, Bloemfontein,9300 South [email protected]

2 Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa

The archosauromorphs of the South African Karoo Basinformed important constituents of the Early to Middle Trias-sic ecosystems following the end-Permian mass extinctionevent. We present data on the osteohistology of the stemarchosaurs Prolacerta, Proterosuchus, Erythrosuchus,Garjania and Euparkeria. Results reveal that the EarlyTriassic non-archosauriform archosauromorph Prolacertaexhibits a poorly defined fibro-lamellar complex, withparallel-fibred bone in some regions. The contemporane-ous Proterosuchus exhibits rapidly forming uninterruptedfibro-lamellar bone early in ontogeny, which becomesslow and cyclical with increasing age. The early MiddleTriassic erythrosuchid Garjania deposited highlyvascularized fibro-lamellar bone interrupted by annuli,whereas the Middle Triassic Erythrosuchus exhibitedhighly vascularized, uninterrupted fibro-lamellar bonethroughout ontogeny. In contrast, the growth of theMiddle Triassic Euparkeria was relatively slow and cyclicalthroughout ontogeny. When our data are combined withthose of previous studies, preliminary results reveal thatEarly and Middle Triassic non-crown group archo-sauromorphs generally exhibit faster growth rates thanmany of those of the Late Triassic. Early rapid growth andrapid attainment of sexual maturity are consistent withlife history predictions for taxa living in the unpredictableconditions following the end-Permian mass extinction.Further research with larger sample sizes will be requiredto determine the nature of the environmental pressureson these basal archosaurs.

Palaeobiology of the Lower Triassic stereospondylLydekkerina (Tetrapoda: Temnospondyli) inferredfrom long bone histology and microanatomy

Canoville, Aurore; Chinsamy-Turan, AnusuyaDepartment of Zoology, University of Cape Town, Rondebosch, Cape Town

[email protected]

Long bone microstructure (histology and microanatomy)provides insight to assess various aspects of tetrapodpalaeobiology, such as ontogeny, growth patterns, andlifestyle. Histology has been extensively applied to thediverse and abundant non-mammalian therapsids fromthe exceptional vertebrate fossil-bearing sequence of the

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Karoo Basin (South Africa), but few studies have beenconducted on amphibians, which were fairly abundant inthe Permo-Triassic terrestrial ecosystems.

Lydekkerina huxleyi, a basal and paedomorphic stereo-spondyl dominated the amphibian fauna from the LowerTriassic Lystrosaurus assemblage Zone. Even though theanatomy of this taxon has been well described, it remainsenigmatic in term of its growth strategies and lifestylehabits.

Earlier studies based on anatomical and taphonomicaldata suggested that the unusually small size of this genus,compared to its Permo-Triassic relatives could be linked toa fast, but shortened development to maintain large andsuccessful breeding populations under difficult environ-mental conditions and may explain the success of theseanimals after the end Permian extinction event. Moreover,Lydekkerina has alternatively been described as aquaticand mostly terrestrial – a controversial hypothesis, sinceTriassic stereospondyls are generally considered to bepredominantly aquatic or semi-aquatic.

In order to document growth patterns and palaeoeco-logical adaptations of this taxon, our study examinedvarious long bones (humerus, radius, femur, tibia, rib) ofseveral specimens of different ontogenetic stages.

Our results show that during early ontogeny, Lydekkerinahad a rapid rate of growth, which slowed down duringlater development. The microanatomy of the long boneswith their thick bone walls and distinctive vacantmedullary cavity suggests that Lydekkerina may have beenamphibious with a tendency to be more terrestrial.

Was Thrinaxodon liorhinus a digger?

Carlson, Kristian1; Fernandez, Vincent2;Abdala, Fernando2; Rubidge, Bruce2; Tafforeau, Paul3

1 Institute for Human Evolution, University of the Witswatersrand, Johannesburg,South [email protected]

2 Bernard Price Institute for Palaeontological Research, University of theWitwatersrand, Johannesburg, South Africa

3 European Synchrotron Radiation Facility, 6 Rue Horowitz Bp 220,38046 Grenoble Cedex, France

Attributing fossorial behaviour to the mammal-likecynodont Thrinaxodon liorhinus has traditionally drawnfrom indirect behavioural evidence. Thrinaxodon speci-mens have been found inside ichnofossils (i.e. burrowcasts) (Damiani et al. 2003) and often fossilize exhibitingcurled-up postures. At stake is whether these distantancestors of mammals may have survived harsh climatesduring the mass extinction event at the Permian-Triassicboundary, approximately 251 million years ago, becauseof their proficiency in creating burrows. Alternatively,Thrinaxodon may have utilized previously constructedburrows, but done so secondarily because they wereincapable of creating new ones, leading to a very differentreconstruction of Thrinaxodon activity patterns and lifehistory. Recent recovery of a complete Thrinaxodon skele-ton inside a burrow cast (Fernandez et al. 2012), plus asecond relatively complete Thrinaxodon skeleton(Damiani et al. 2003), provide two sets of associatedpostcrania for analysis. Using synchrotron scanning fromthe European Synchrotron Radiation Facility (Grenoble,

France), virtual renderings of postcranial elements wereproduced. Structural comparisons between Thrinaxodonand extant wombats (Genera Vombatus and Lasiorhinus)were performed to identify potential fossorial adaptationsin limbs of the former. Wombats are well-known fossorialanimals roughly equivalent in body size to Thrinaxodon.Such comparisons permit assessment of direct structuralevidence for fossorial adapations in Thrinaxodon liorhinus,or determine whether their digging ability has been inac-curately attributed because of guilt by association.

Damiani R, Modesto S, Yates A. & Neveling J. 2003. Earliest evidence ofcynodont burrowing. Proceedings of the Royal Society of London B 270,1747–1751.

Fernandez, V., Abdala, F., Carlson, K.J., Collins Cook, D., Rubidge, B.,Yates, A. & Tafforeau, P. 2012. Synchrotron radiation sheds light onmammal forerunners entombed in their burrow. Palaeontologia africana47: 35.

Bone microstructure of polar and temperateEdmontosaurus

Chinsamy, Anusuya1; Thomas, Daniel B.1;Tumarkin-Deratzian, Allison R.2; Fiorillo, Anthony R.3

1 University of Cape Town, Zoology Department, Rondebosch, South [email protected]

2 Dallas Museum of Natural History, Dallas, TX 75315, U.S.A.3 Temple University, Department of Earth and Environmental Science, Philadelphia,

Pennsylvania 19122, [email protected] / [email protected]

Dinosaurs have a worldwide distribution, and appear tohave inhabited a range of latitudinal zones, includinghigh latitudinal localities that were well within the Arcticand Antarctic regions, i.e. areas today known to havedramatic changes in temperature and light during thecourse of a single year. The Upper Cretaceous PrinceCreek Formation of the North Slope of Alaska is one of therichest high latitudinal dinosaur-bearing deposits. Skeletalmaterial and track assemblages of a wide diversity ofnonavian dinosaurs (e.g. large and small bodied theropods,ceratopsians, pachycephalosaurs, hypsilophodontids,and hadrosaurs) have been recovered from this locality.One of the major questions is whether these high-latitudedinosaurs overwintered or migrated during the winters.

Since it is well documented that various aspects of thebiology and life history of extinct vertebrates is recordedin the microstructure of their fossilized bones, we com-pared the long bone histology of Edmontosaurus from thePrince Creek Formation (69.5 ± 1.5 Ma) with that ofEdmontosaurus from the more temperate, HorseshoeCanyon Formation (70.0 ± 3.0 Ma) of Canada. Our samplecomprised seven femora, four humeri and two tibiae fromthe polar Edmontosaurus, and three femora and one tibiafrom temperate Edmontosaurus.

We found that the bone microstructure of the polarEdmontosaurus exhibited highly vascularized, fibro-lamellar bone with periodic changes in channel organiza-tion i.e. from a reticular oriented ’vascularization’ to amore circumferential orientation. In contrast, such alter-nating changes in the vascularization were not consis-tently observed in the bones of Edmontosaurus from themore temperate Canadian locality.

The differences observed in the microstructure ofEdmontosaurus from the high and low latitude localities,

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suggests that they grew differently. From the type of bonemicrostructure evident, we deduce that polar Edmontosaurusendured the long winter night, while temperate Edmonto-saurus lived in a perennially moderate environment.Thus, we propose that the regular alternating vascu-larization pattern observed in the polar dinosaurs is anoverwintering signal in response to an annual change inthe prevailing environmental conditions.

A new pareiasaur reptile from the TapinocephalusAssemblage Zone, middle Permian of the Karoo

Cisneros, Juan Carlos1; Rubidge, Bruce2

1 Centro de Ciências da Natureza, Universidade Federal do Piauí, Teresina, [email protected]

2 Bernard Price Institute for Palaeontological Research, University of theWitwatersrand, Johannesburg, South [email protected]

Pareiasaurs are large, heavily-built members of theParareptilia, that had a global distribution and constitutedimportant components of the herbivore niche in terres-trial faunas during middle to late Permian times. Wereport a new pareiasaur species from the middle Permianof South Africa. The fossil was found in the Prince AlbertDistrict, Western Cape, in the Abrahamskal Formation,upper Tapinocephalus Assemblage Zone (AZ). The speci-men consists of a fragmentary cranium with mandible inocclusion, and an articulated postcranium including mostof the vertebral column, ribs, pectoral and pelvic girdles,and some long bones. Despite being very damaged, theskull of the new specimen has a well-preserved marginaldentition that allows direct comparisons with mostpareiasaurs known in the Karoo and elsewhere. The newfossil is easily distinguished from other pareiasaurs byhaving only five cusps on each upper marginal tooth,being the lowest number of dental cusps in any knownspecies of pareiasaur. Other pareiasaurs have upper toothcusp numbers ranging from seven (Bradysaurus spp. andNochelesaurus alexanderi) to fifteen (Anthodon serrarius).Another notable feature in the teeth of the new specimenis their shape, having three closely spaced cusps – second,third and fourth – that rise well beyond the level of thefirst and fifth cusp, providing altogether a rather elon-gated outline to each marginal tooth that contrasts with awider outline in the teeth of most pareiasaur species.These characteristics allow us to recognize a new speciesof pareiasaur in the Tapinocephalus AZ. The extreme lownumber of dental cusps present on this pareiasaur is remi-niscent of the condition in the nycteroleters, a group ofparareptiles that is well represented in the MiddlePermian of the Russian Platform, which is considered toconstitute the sister group of the Pareiasauria. In mostnycteroleters the marginal teeth are thin, elongated andmonocuspid, and in the genera Bashkyroleter and Riphaeo-saurus, three cusps are present. The teeth of the newpareiasaur, thus, constitute an intermediary conditionbetween the elongated, mono- or tri-cuspid teeth ofthe nycteroleters and the wide teeth, with several cups,present in all other pareiasaurs. In this way, the teeth of thenew pareiasaur fill a gap in tooth evolution towardsherbivory in the Nycteroleter-Pareiasaur clade of theParareptilia.

Actualistic investigation of bone modification bycaracal (Caracal caracal) and honey badger(Mellivora capensis) on domestic rabbit carcasses

Cohen, Brigette; Kibii, JobInstitute for Human Evolution, University of the Witwatersrand, Private Bag 3,

WITS 2050, Johannesburg, South [email protected] / [email protected]

The involvement of carnivores and birds of prey in boneaccumulation in archaeological and palaeontologicalfossil assemblages has often been inferred through diag-nostic bone modification. The bulk of actualistic and fieldwork investigation, has concentrated on assessing thenature of bone modifications by large carnivores on largebodied mammals, much to the neglect of smaller preda-tors and prey. This study investigated the nature of bonemodification of mesomammal (leporid) carcasses bycaptive small carnivores including the caracal (Caracalcaracal) and the hitherto taphonomically undocumentedhoney badger (Mellivora capensis). Domestic rabbitcarcasses were fed to the two captive carnivore species atthe Johannesburg Zoo. Scatological remains werecollected, the bones removed, cleaned and taphonomi-cally assessed. Preliminary analysis indicates that thehoney badger is highly destructive of rabbit remains. Atotal of 392 identifiable bone fragments were recoveredfrom the honey badger scats, representing only 16% of therecovered bones. Average bone length was 10.3 mm andthe maximum length recovered was 28.1 mm. The assem-blage was made up primarily of vertebral fragments,particularly from the lumbar region, and rib fragments.Cranial and femoral remains are also high as were phalan-ges. There was a total of 237 tooth marks in the assem-blage, the majority made up of crush marks and scours.The majority of bones showed evidence of digestive dam-age. A template deriving from actualistic investigation iscrucial in the assessment of fossil assemblages, particu-larly in determining taphonomic agents and processes in-volved in the accumulation of mesomammals in fossilassemblages.

Corked! The history and importance ofhumanities’ favourite beverage

Cohen, Brigette; Val, AuroreInstitute for Human Evolution, University of the Witwatersrand, Private Bag 3,

WITS 2050, Johannesburg, South [email protected] / [email protected]

It was common practice 20 years ago for archaeologiststo clean out the sediments contained in any recoveredcontainers or jars and to analyse only the potsherds them-selves. However, recent research by such experts as PatrickMcGovern has shown the importance in preserving andanalysing the surrounding sediments, a principle that hasalso been successfully applied to palaeontological excava-tions. Today it has become just as important, if not supe-rior, to analyse residues found within jars and containers.Indeed the contents of sealed containers can last surprisinglengths of time under the right conditions; in 2011 soupwas recovered from 2000-year old bronze and clay con-tainers in China (Hilgers 2012). Thus this research inconjunction with ancient engravings and texts gave rise tothe field of ‘dipsology’. Chemical analysis of potsherds

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has revealed the oldest beers, wines and grogs (Finally ause for them!). Not only that but independent brewershave got on the experimental archaeology bandwagonattempting to recreate such forgotten masterpieces (geeand I thought archaeologists were nerdy!). Dipsologyprovides us with more than just a peek into how ourfavorite beverage may have looked or tasted or even whatit was made from. It provides an understanding of whendomestic fermentation began and even indicates that itmay have performed a fundamental role in the creation ofthe modern static civilization. What if this behaviour wentback even further? Recent archaeological research inSouth Africa has been pushing back the boundaries ofearliest modern human behaviour with the recovery ofetchings, jewellery and even paint containers. Perhapsone of the most important modern human behavioursalso began in Africa.

Hilgers, L. 2012. Ancient Chinese takeout. Archaeology Magazine, January/February.

Middle Permian biodiversity in the Karoo: abiostratigraphic review of the TapinocephalusAssemblage Zone

Day, Mike; Rubidge, Bruce; Güven, SaniyeBernard Price Institute for Palaeontological Research, School of Geosciences,

University of the Witwatersrand, Johannesburg, South [email protected] / [email protected] /[email protected]

The Beaufort Group of South Africa contains one of themost complete Permo-Triassic tetrapod fossil records inthe world. Its lower strata are of late Middle Permian age,comprising up to 2700 m of sediment at its thickest pointalong the Cape Fold Belt. These sediments fall withinthree faunal assemblage zones of which the second, theTapinocephalus Zone, is stratigraphically the thickest andhas the highest diversity of fossil genera (Rubidge 1995).In addition, the fauna in the Tapinocephalus Zone is domi-nated by the Dinocephalia, a group which was highlysuccessful towards the end of this stage but which becameextinct before the beginning of the Late Permian. A linkbetween the disappearance of the Dinocephalia and amarine extinction in China has already been proposed(Retallack et al. 2006), but there is no direct evidence tosupport this. An understanding of the TapinocephalusZone is therefore crucial to understanding Middle Perm-ian tetrapod diversity, while the transition into the suc-ceeding Pristerognathus Zone provides the opportunity tostudy the extinction of the Dinocephalia.

The outcrop area of the Tapinocephalus Zone extendsfrom the Eastern Cape near East London, west toLaingsburg in the Western Cape and northwest to VictoriaWest in the Northern Cape Province. Over such a widearea, the stratigraphy varies laterally as well as thinningrapidly towards the north. The strata in the south are alsoheavily folded and together these have made it difficult toconduct accurate biostratigraphic studies. Utilization ofthe Beaufort Vertebrate Database housed at the BPI com-bined with satellite imagery, extensive fieldwork andstratigraphic collecting conducted over many years, hasenabled examination of the the Tapinocephalus Zone at afar higher biostratigraphic resolution. Stratigraphic

ranges of vertebrate taxa are more distinct from each otherthan is currently understood with the ranges of mostgenera restricted to the upper part of the zone. Dino-cephalians also extend higher in the stratigraphic succes-sion than previously documented.

Retallack, G.J., Metzger C.A. et al. 2006. Middle-Late Permian massextinction on land. Geological Society of America Bulletin 118(11-12),1398–1411.

Rubidge, B.S. (ed.) 1995. Biostratigraphy of the Beaufort Group (KarooSupergroup). South African Committee for Stratigraphy Biostrati-graphic Series.

Early diagenesis in Late Jurassic sauropod teethrevealed by transmission electron microscopy

Dumont, Maïtena1; Kostka, Aleksander1; Tütken, Thomas2

1 Abteilung für Werkstoffdiagnostik und Technologie der Stähle, Max-Planck-Institutfür Eisenforschung, Düsseldorf, [email protected]

2 Steinmann Institute of Geology, Mineralogy and Palaeontology, University ofBonn, Germany

The stable isotope compositions (Ç13C, Ç18O, 13C–18O) of

dinosaur tooth enamel is widely used to study the diet,thermophysiology and paleoenvironment of the dino-saurs. It is then important to determine if original isotopecompositions are still preserved in the fossil teeth and tounderstand possible geochemical and mineralogical mod-ifications during the fossilization process. For this purposethe microstructure of dental tissues of teeth from large Ju-rassic sauropods such as Branchiosaurus brancai from theTendaguru beds in Tanzania (Africa) and Camarasaurus sp.from the Morrison Formation (U.S.A.) were investigatedusing transmission electron microscopy (TEM). Recently,clumped isotope thermometry of the enamel bioapatitefrom the same teeth determined physiologically plausi-ble, mammalian-like body temperatures for the sauro-pods that suggest good preservation of these teeth andtheir stable isotope compositions (Eagle et al. 2011).

Our TEM results show different stages of diageneticalteration between the investigated tooth samples. TheTendaguru teeth are well preserved compared to the Mor-rison material, confirming previous conclusions drawn bystable and clumped isotope compositions (Eagle et al.2011).

Furthermore, enamel is generally less altered bydiagenesis than the dentine of the same teeth. Indeed, thecrystal size in the enamel does not change drasticallybetween altered and well-preserved tooth areas, whereasan increase of the dentine crystals in size and in theirwidth/thickness is observed. TEM results coupled withEDX (energy dispersive X-ray) analysis of the chemicalcomposition and diffraction pattern allow us to discernprecisely changes in the apatite crystal structure duringdiagenesis. Secondary mineral infillings, when presentin the dentine tubuli, are also analysed and identifiedprecisely by indexing their diffraction patterns.

This approach enables to characterize the history of thediagenetic alteration in teeth of different taphonomicsettings.

Eagle, R.A., Tütken, T., Martin, T.S., Tripati, A.K., Fricke, H.C., Connely,M., Cifelli, R.L. & Eilen, J.M. 2011. Dinosaur body temperatures deter-mined from isotopic (13C–18O) ordering in fossil biominerals. Science333(6041), 443–445.

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The impact of mining on the Cradle of humankindWorld Heritage Site

Durand, FrancoisDepartment of Zoology, University of Johannesburg

[email protected]

South Africa has an exceptional cultural and naturalheritage; the largest astrobleme in the world, the largestlayered igneous intrusion in the world containing some ofthe richest ore deposits in the world, the largest goldrepository in the world, some of the oldest fossils in theworld, the largest terrestrial Permo-Triassic sequence inthe world containing the most synapsids and some of theoldest dinosaurs in the world, the most hominin fossils,spanning the largest part of human prehistory, in theworld and an unparalleled diversity of fauna and flora.This geological, palaeontological, archaeological and eco-logical wealth has spawned two industries – mining andtourism which are in conflict with each other. Conserva-tion of our natural and cultural resources is threatened byshort-term opportunism and mismanagement. It isproposed that this conflict arises from the fact that theconnections between the geological, ecological and cul-tural wealth are not appreciated nor understood. In therecent past the threat of acid mine drainage (AMD) fromthe mine void adjacent to the Cradle of HumankindWorld Heritage Site (COHWHS) was ignored. The influxof megalitres of AMD into the COHWHS in 2011 served asa wakeup call to everybody. The climate change led tohigher rainfall in the region which resulted in the eleva-tion of the water table and an increase in the efflux ofAMD from the mine void on the West Rand. This can beseen in the rise of the level of the ’underground lake’ inSterkfontein Caves and the Rietspruit which has turnedinto a perennial river carrying AMD at a pH below 3. TheRietspruit is a tributary of the Bloubankspruit which flowsthrough the southern part of the COHWHS past severalpalaeontological sites. The groundwater in the region issimilarly affected as a plume of AMD is spreading fromthe mine void into the Swartkrans dolomitic compart-ment underlying the southern part of the COHWHS.

Synchrotron radiation sheds light on mammalforerunners entombed in their burrow

Fernandez, Vincent1; Abdala, Fernando1;Carlson, Kristian J.2; Collins Cook, Della3;Rubidge, Bruce1; Yates, Adam1; Tafforeau, Paul4

1 Bernard Price Institute for Palaeontological Research, University of theWitwatersrand, South [email protected] / [email protected] /[email protected] / [email protected]

2 Institute for Human Evolution, University of the Witswatersrand, South [email protected]

3 Department of Anthropology, Indiana University, [email protected]

4 European Synchrotron Radiation Facility, [email protected]

The Karoo Basin of southern Africa has provided numer-ous examples of vertebrate burrowing behaviours in strataspanning the Permo-Triassic boundary (PT, ~251 Myr).The use of underground shelters has been interpreted asevidence of their importance in providing a stable envi-

ronment that protected occupants against harsh condi-tions during the PT mass extinction event. During the lastdecade, several burrow casts with occupants have beendiscovered; some include aggregations of several individ-uals or even different species. A key observation was thefact that several of these specimens were therapsids, thevery distant ancestors of mammals. Fifty per cent of modernmammals are capable of burrowing; this is viewed as anancient behavioural trait that pre-conditioned burrowersurvival under the harsh environments and successiveglobal Permo-Triassic extinctions.

A major limitation with such specimens is the fact thatthe fossils are enclosed in burrow casts and thereforeinvisible without preparation. However, classic prepara-tion (physical or chemical removal of the surroundingrock) destroys all information about the sediment infill.Moreover, observations on prepared skeletal remains areoften limited to a single side because it is necessary toleave intact matrix in order to maintain the integrity ofspecimens.

In the last few years, third generation synchrotron radia-tion-based X-ray microtomography (SR- CT) has providednew ways to increase the quality of data obtained fromfossil specimens. Synchrotrons allow use of other methodsthat are thousands of times more sensitive and utilizephase-contrast effects, such as propagation phase contrastmicrotomography (PPC-SR-µCT). While many synchro-tron X-ray imaging beamlines are interested in develop-ing narrow X-ray beams, the European SynchrotronRadiation Facility (ESRF) offers the possibility to scanlarge specimens and keeps improving the possibilities fornon-destructive investigations of fossils.

Here we present results obtained using the large X-raybeam of the ID17 beamline of the ESRF on fossilized tetra-pods preserved in burrow-casts from the Early Triassic ofthe Karoo Basin. Images produced by PPC-SR CT exhibita high degree of contrast and accuracy. Whole burrowswere imaged with a voxel size of 45 µm, revealing everystructure of the skeletons as well as fine structure of thesediment infill. The images and resultant rendering allowus to understand the thanatocoenose or death assemblageand observe information about the behaviour of mammalforerunners.

Distribution patterns of Devoniantetrapodomorphs revealed by studyof pectoral girdle morphologies

Gess, RobertBernard Price Institute for Palaeontological Research, University of the

Witwatersrand, South [email protected]

Sarcopterygians include ’fish-like’ sarcopterygians (lobefinned fishes) and tetrapods. Intermediate forms, looselygrouped as panderichyids or elpistostegids, provide cluesregarding arguably the most important transition in ver-tebrate evolution, that from finned fish to digit-bearingtetrapods. The most important morphological changes in-clude those affecting the shoulder girdle, which permittedcritical changes in paired appendage use.

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The pectoral girdle of sarcopterygians includes anendoskeletal scapulocoracoid and a number of dermalelements, most importantly the cleithrum and clavicle. In’fish-like’ sarcopterygians these bones and other dermalshoulder girdle bones (which may include an anocleithrum,extracleithrum, supracleithrum and interclavicle) form arigid dermal skeleton articulating with the skull. Thescapulocoracoid is attached to the medial face of thecleithrum and sometimes also the clavicle. It has a posteri-orly directed glenoid for articulation of the pectoral fin,which does not protrude far beyond the posterior marginof the dermal girdle.

Tristichoperid fish, the most derived ’fish-like’ sarco-pterygians, in terms of tetrapod evolution, includeEusthenopteron. The two halves of the dermal pectoralgirdle of Eusthenopteron are linked by a thin interclavicle,on either side of which the girdle includes a clavicle,cleithrum, anocleithrum and supracleithrum. Thecleithrum dominates the dermal girdle and is subrect-angular and upright (in lateral view) with a ventromedialextension at the attachment point of the clavicle. Thescapulocoracoid is attached to the medial wall of theventrolateral angle of the cleithrum.

The cleithrum of the elpistostegids Panderichthys andTiktaalic is more dorso-posteriorly angled and lacks aventromedial extension. It is ventrolaterally reduced toallow extensive exposure of the scapulocoracoid. This ismuch larger than that of Eusthenopteron, has extensivecontact with the clavicle and a more laterally orientatedarticulation for the pectoral appendage. Greater exposureof the scapulocoracoid permitted the fin not only to assumepostures suitable for swimming but also for a substratesupported upright stance.

In Acanthostega, the best known early tetrapod (withdigits), the pectoral girdle is detached from the back ofthe skull. The cleithrum is unornamented and far morereduced posteroventrally, further exposing the scapulo-coracoid. In anterior view a distinct postbranchial laminaforms an anteromedially directed flange. A ridge extendsposteroventrally from the cleithrum onto the scapulo-coracoid.

Rapid morphological changes spanning the emergenceof tetrapods give cleithra diagnostic value in the study ofdisarticulated Devonian sarcopterygians. More than onegrade of ’advanced’ sarcopterygian is thereby identifiedfrom the Late Devonian Waterloo Farm locality, withimportant implications for the range of environmentalsettings in which transitional forms occurred.

Palynofacies patterns of the Northern WitbankBasin (South Africa): clue to decipher Permianpalaeoenvironmental and climate change

Götz, Annette E.1; Ruckwied, Katrin2

1 Department of Geology, Rhodes University, South [email protected]

2 Shell International Exploration and Production B.V., the Netherlands

The Main Karoo Basin hosts an important coal resourceof South Africa, the Witbank coal field, which comprisesthe basin’s northeasternmost economic coal deposits.

These sediments are part of the Ecca Group and weredeposited subsequent to the melting of the Dwyka ice.One of the most important coal seams of the WitbankBasin, the No. 2 coal seam, represents postglacial fluvio-deltaic deposits and here, we report on new palynologicaldata from a highly proximal setting of the northern basinmargin.

The sedimentary series includes coarse-grained topebbly-sandstones, with an abrupt upward transitioninto fine-grained sediments and coal, through cross-stratified medium- to coarse-grained sandstones, and hor-izontally-laminated fine- to medium-grained sandstonesand siltstones. The highly volatile bituminous coals arecharacterized by significant amounts of inertinite and lowmaturity, with a vitrinite reflectance ranging from 0.62%to 0.8%, and medium to high ash content, ranging be-tween 7.8 wt% and 73.6 wt%. The sedimentary organicmatter content clearly documents changes in the paly-nomorph assemblage and variations in the amount andthe type, size and shape of plant debris. Generally, thepalynofacies are characterized by high proportions ofopaque phytoclasts. Amorphous organic matter is charac-teristic of laminated siltstones and coals. The palynologicalrecord indicates a cold climate fern wetland communitycharacteristic of lowland alluvial plains with an uplandconifer community in the lower part of the coal seam.In the upper section, these communities are replaced bya cool-temperate cycad-like lowland vegetation andgymnospermous upland flora.

Our data establish a base for further integratedpalynological-palaeobotanical studies on terrestrial plantvegetation and its response to climate amelioration. Thecyclic patterns recognized in the sedimentary series westudied will, furthermore, allow interpretation ofpalaeoenvironmental change at greater time resolution,which will lead to a better understanding of the timing offloral change.

Five million-year-old Cetacean population fromLangebaanweg, west coast of South Africa

Govender, Romala1; Chinsamy-Turan, Anusuya2

1 Natural History Department, Iziko South African Museum, Cape [email protected]

2 Department of Zoology, University of Cape Town, [email protected]

Langebaanweg (LBW) is a world renowned and prolificAfrican Tertiary fossil site. It records local Mio-Pliocenethe fauna, flora and environment. Among this treasuretrove of fossil vertebrates are the remains of marinemammals. Our improved understanding of the terrestrialvertebrate fauna and palaeoenvironment has highlighteda lack of knowledge of the marine context. We report hereon the preliminary results of my analysis of the cetaceanremains from LBW’s ‘E’ Quarry.

The condition of the fossils ranges from fragmentary towell preserved; the latter, in particular, being bones of thepetrotympanic region. Our study focusses on these bonesas they are reasonably readily identified, at least to genericlevel. The ear bones are predominantly of mystecetes

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and include balaeonpterid and balaenid forms. Fivemillion years ago a lagoon or estuarine system existed atLangebaanweg. Hypothtically the occurrence of cetaceanremains appears to be the result of their larger size and‘floatability’, which allowed carcasses to be beached;thereafter, storm surges refloated partially skeletonizedcarcasses, which further disintegrated and fragmentedbefore being moved upstream where some came to rest inthe channels or scour pools in which they were found.

Taxonomy of tapinocephalid Dinocephalia fromthe South African Karoo Basin

Güven, Saniye; Rubdige, Bruce; Abdala, FernandoBernard Price Institute for Palaeontological Research, Palaeosciences Centre,

University of the Witwatersrand, South [email protected] / [email protected] /[email protected]

Tapinocephalids (Therapsida: Dinocephalia) are animportant component of the Middle Permian terrestrialecosystem. They are known from South Africa, Zimbabwe,Tanzania, Russia, and Brazil. South Africa contains by farthe most abundant and morphologically varied fauna ofthis group which are the most important herbivorouscomponent from the fauna of the Tapinocephalus Assem-blage Zone. Recent studies indicate a large number oftapinocephalid genera compared to other herbivoroustherapsids in any particular biozone of the BeaufortGroup. Additionally, most tapinocephalian taxa are basedon either fragmentary or heavily weathered types,suggesting the need for taxonomic re-examination. As aresult of our taxonomic survey we reduced the recognizedgenera of tapinocephalids from 18 to 9. This revision indi-cates that the diversity of tapinocephalid dinocephaliansin the Tapinocephalus AZ was overestimated. This willmodify our understanding of the Middle Permian extinc-tion event that appears to have affected dinocephalians inparticular.

First tapinocephalid dinocephalian from thePristerognathus Assemblage Zone (KarooSupergroup, South Africa), including newinformation on Criocephalosaurus(Therapsida: Dinocephalia)(Poster)

Güven, Saniye1; Day, Michael1; Almond, John E.2;Abdala, Fernando1; Rubidge, Bruce S.1

1 Bernard Price Institute for Palaeontological Research, Palaeosciences Centre,University of the Witwatersrand, South [email protected] / [email protected] /[email protected] / [email protected]

2 Natura Viva CC, P.O. Box 12410 Mill Street, Cape Town, 8010 South [email protected]

Tapinocephalid dinocephalians were impressively largeherbivores from the Middle Permian of South Africa andare considered to be stratigraphically restricted to theEodicynodon and Tapinocephalus Assemblage Zones of theKaroo basin. We report the first tapinocephalid from thelower Pristerognathus Assemblage Zone. The specimen,which comprises the posterior portion of the skull, is iden-tified as Criocephalosaurus (Criocephalus Broom, 1928[nomen nudum]). This poorly-known genus comprises twospecies: C. vanderbyli represented by a weathered incom-

plete posterior cranial roof (holotype) and some cranialfragments; and C. gunyankaensis based on posterior cranialroof fragments. The first species is known from strata ofthe Abrahamskraal Formation, South Africa andstratigraphically occurs in the Tapinocephalus AssemblageZone; whereas C. gunyankaensis was discovered inGunyankas Kraal, Zimbabwe, from horizons consideredto be equivalent to the Tapinocephalus Assemblage Zone ofSouth Africa. Due to the fragmentary and poorly pre-served nature of the specimens this genus has notreceived much attention. The issue is further complicatedby the fact that the holotype of C. gunyankaensis is missing.This specimen was described in a very short paper with-out any diagnosis. Accordingly the only reliable morpho-logical data of the genus is based on the genotype. Inaddition to the new fossil from the Pristerognathus Assem-blage Zone, a specimen referable to Criocephalosaurus hasbeen discovered in the palaeontological collection of theNatural History Museum, London. Both specimens allowrevision of the cranial description of Criocephalosaurus andhave facilitated additional characters which are not pres-ent in the holotype. This discovery raises the possibility ofthe unexpected survival of dinocephalians beyond theTapinocephalus Assemblage Zone age. It also highlights thenecessity for more-detailed palaeontological sampling inthe lower levels of the Pristerognathus Assemblage Zone ofthe Karoo Basin.

Mosasaur long bone microanatomical andhistological features

Houssaye, Alexandra1; Lindgren, Johan2

1 Steinmann Institute of Geology, Mineralogy and Palaeontology, University ofBonn, [email protected]

2 Department of Geology, GeoBiosphere Science Centre, Lund University, Sweden

Mosasauroids are an extinct group of medium-sized togiant squamates that succeeded in establishing them-selves as the dominant marine reptiles in a brief 27 millionyears at the end of the Cretaceous. Osteohistologicalstudies of mosasauroids have hitherto been limited to ribsand vertebrae. Here we present data on long bonemicroanatomical and histological features of varioushydropelvic (i.e. with no sacrum) forms, including theplesiopedal (i.e. with terrestrial-like limbs) Dallasaurusand hydropedal (i.e. with paddle-like limbs) Angolasaurus,Globidens, Halisaurus, Mosasaurus, Platecarpus, Plotosaurus,and Prognathodon. In all hydropedal genera, the bonemicroanatomy is characterized by a spongious organiza-tion with no medullary cavity; a condition which isconspicuously different from that in Dallasaurus. Thehistology shows a dominance of lamellar-zonal bone(although fibrous bone can occur locally), and a well-developed vascular network with a predominantly longi-tudinal orientation. Comparisons are made with extantterrestrial and semi-aquatic lizards, such as Amblyrhynchus,Tupinambis, and Varanus, as well as obligate aquatic tetra-pods, such as ichthyosaurs, plesiosaurs and cetaceans.Histological features suggest relatively low metabolicrates in mosasaurs, where high body temperaturespresumably were maintained by gigantothermy.

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Do body size trends in nonmammaliantherapsids reveal the influence of theend-Permian extinction?

Huttenlocker, Adam1; Sidor, Christian2;Botha-Brink, Jennifer3

1 Department of Biology, University of Washington, [email protected]

2 Burke Museum and Department of Biology, University of Washington,Seattle, Washington 98195, U.S.A.

3 Karoo Palaeontology, National Museum, Bloemfontein, 9300 South Africa

Post-extinction body size reductions in animals (or’Lilliput phenomena’) represent a recently identifiedfeature of mass extinctions. Such have been identified inearliest Triassic marine invertebrates and anecdotally inSouth African temnospondyls and nonmammaliantherapsids following the end-Permian extinction (ca. 252Ma). However, field observations of post-extinction sizereductions in terrestrial vertebrate communities have notpreviously been subjected to rigorous statistical testing.Here, we characterize body size trends in Permian andearly-to-mid-Triassic eutheriodonts, an ecologically variedtherapsid subclade which outnumbered contemporarydicynodont herbivores in terms of species richness duringthe earliest Triassic and eventually gave rise to mammals.This research addresses the simple question: Are bodysize decreases observed among and/or within eutherio-dont taxa following the end-Permian extinction?

Preliminary analysis on a database of measurementsfrom more than 343 eutheriodont specimens indicateslarge body sizes and increasing size disparity in PermianCistecephalus and Dicynodon Assemblage Zones, followedby a slight decrease in size disparity and reduced overallbody size in the Triassic Lystrosaurus Assemblage Zone.Basal skull length (BSL) as well as femur and humerusmidshaft diameter were used as relative proxies for sizeas (1) complete skeletons were not always available forprecise mass estimates and (2) their proportions share anapproximate relationship with body mass. Our findingscorroborate earlier results on BSL in the Permo-Triassic(P-Tr) therocephalian Moschorhinus, which suggestedsignificant within-lineage cranial size reductions in P-Trsurvivor lineages. More general trends corroborate signif-icant decreases in BSL and limb bone dimensions ineutheriodonts as a whole. Finally, to correct for theimpacts of longer term phylogenetic trends, it is necessaryto recognize potential phylogenetic constraints onobserved body size distributions. In other words, areobserved body size trends stochastic or ’driven’ and, ifdriven, are they explainable by extrinsic processes actingon body size distributions during the P-Tr transition? Wewill explore statistical approaches which include (1)regression on dissimilarity matrices (Mantel tests) and (2)model-based tree randomization procedures in order toidentify the extent to which size distributions are explain-able by phylogenetic distance and tree structure. Our nullprediction is that size shifts are stochastic and largely re-flect tree structure (i.e. closely related clades should bemore similar in size than phylogenetically disparateclades). However, initial results from Mantel tests indicatethat size disparity correlates poorly with pairwise phylo-genetic distance within the therocephalian subclade.

Alternatively, if extrinsic factors indiscriminately acceler-ated rates of life history evolution across clades, thenobserved size distributions should show significantanti-signal in earliest Triassic clades (a signature of adap-tive radiations).

Morphological variation of Sterkfontein andSwartkrans first metatarsals by linear featureextraction on morphometric colour maps

Jashashvili, Tea1; Dowdeswell, Mark R.2;Carlson, Kristian J.1; Marchi, Damiano1;Nshimirimana, Robert3

1 Institute for Human Evolution, University of the Witwatersrand, South [email protected]

2 School of Statistics and Actuarial Science, University of the Witwatersrand,South [email protected]

3 NECSA, Radiation Sciences Department, Pretoria, South Africa

Four first metatarsals, two from Sterkfontein (StW562,StW595) and two from Swartkrans (SKX5017, SK1813) areanalysed. SKX5017 is attributed to Paranthropus robustus.Taxonomical attribution of SK1813 is unsure althoughmorphological comparisons show similarity with SKX5017.StW562 and StW595 are from Member 4 deposits at Sterk-fontein and are described as intermediate in morphologybetween modern humans and apes. Unfortunately, theyhave yet to be attributed to a genus or species. The mosaicnature of morphological trait distribution on the metatar-sals complicates functional interpretations: early homininswere unquestionably bipedal but possessed a uniquetoe-off mechanism different to modern humans. This pa-per shows preliminary results of first metatarsal shaft cor-tical thickness distribution in living hominoids. Thisinformation is used to provide a better interpretation ofthe bending and compressive stress patterns in early aus-tralopithecines.

First metatarsals of chimpanzees, gorillas and modernhumans are compared to those of fossils. For each meta-tarsal we took 17 cross-sections from 25–65% of the shaftmechanical length. Distance between consecutive crosssections is 2.5% of metatarsal mechanical length. Wemeasured cortical thickness at different radii of each crosssection at 1° increment. Cortical thicknesses are standard-ized by size and normalized between 0 and 1. Distributionof thicknesses along shafts is shown using colour maps.We subsequently perform Linear Feature Extraction onmorphometric colour maps of the shafts in order to showthe differences between presented taxa.

Results show that chimpanzees and gorillas differ frommodern humans in that great apes have much thickercortical bone distribution distally on the shaft, dorso-lateral thicker cortical shafts proximally, and dorso-plantar thicker cortical bone distally. Modern humans donot show much variation along the shaft with predomi-nantly thicker cross sections along the dorso-plantardirection. Both Swartkrans (SKX5017, SK1813) andSterkfontein (StW562) metatarsals have similar distribu-tion patterns to chimpanzees and gorillas. StW959 fromSterkfontein has a much more human-like distribution.We confirm that StW562 represents the ’robust’ form ofhominin fossil from Sterkfontein Member 4. StW959

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represents the ’gracile’ form (Au. africanus) fromSterkfontein, and has modern human characteristics inthe distribution of cortical thickness that could be associ-ated to a similar shaft bending regime.

Sediment sources of the aeolian ClarensFormation, Karoo Supergroup

Jinnah, Zubair1; Roberts, Eric2; Dirks, Paul2

1 School of Geosciences, University of the Witwatersrand, Johannesburg,South [email protected]

2 School of Earth and Environmental Sciences, James Cook University,Townsville, [email protected] / [email protected]

Detrital zircon geochronology is a widely used tech-nique in sedimentary basin analysis and has becomeimportant in finding maximum depositional ages andsource areas for otherwise poorly constrained strati-graphic units. The aeolian Clarens Formation, which is theuppermost sedimentary unit in the Karoo Supergroup, isa prime locality for detrital zircon analysis becauseClarens sediment is derived largely from reworking ofsediment from older Karoo units. Zircons in the ClarensFormation thus contain populations from all significantsources of the Karoo Basin.

The Clarens Formation was sampled in the northwest(Free State), south (Eastern Cape) and east (KwaZulu-Natal) of the extent of outcrop. Zircon populations aredominated by groups originally derived from Pan-Africanage (approximately 700–500 Ma) and Namaqua-Natal age(approximately 1 Ga). A small population of Archaeanzircons are also present in the Clarens Formation.Phanerozoic zircons include populations from the Ordo-vician, Silurian, Permian and Triassic. Ordovician andSilurian grains are potentially recycled from the CapeSupergroup, and may be ultimately derived fromAntarctica or South America, whereas Permian zirconsmay potentially be derived from reworked ash beds in thelower parts (Ecca and Beaufort Groups) of the KarooSupergroup. One Triassic grain, with an age of 212.7 +5.6 Ma, is enigmatic as it is much younger than any ashdocumented from the Karoo Supergroup, and does notoverlap temporally with any kimberlites in the region.The exact source of this grain remains uncertain but itspresence highlights the potential of detrital zircons toprovide age constraint for the Triassic-Jurassic fill of theKaroo Basin with continued detrital zircon geochronol-ogy.

Stratigraphy and sedimentology of theTapinocephalus Assemblage Zone(Abrahamskraal Formation) in the areaaround Merweville, South Africa

Jirah, Sifelani; Rubidge, BruceBernard Price Institute for Palaeontological Research, School of Geosciences,

University of the Witwatersrand, Johannesburg, South [email protected] / [email protected]

By far the thickest biozone of the Beaufort Group is theTapinocephalus Assemblage Zone, which, despite its thick-ness and the presence of tetrapod fossils, has not yet beenbiostratigraphically subdivided in a satisfactory way.Rossouw & De Villiers (1953) proposed a threefold subdi-

vision of the Tapinocephalus Biozone and Boonstra (1969)confirmed this subdivision. Keyser & Smith (1978) definethe upper limit of the Tapinocephalus Biozone by the lastoccurrence of dinocephalians which they say coincideswith an extensive green ’chert’ band about 120 m belowthe Poortjie Sandstone.

Recent stratigraphic and sedimentological analysis ofthe Tapinocephalus Assemblage Zone rocks aroundMerweville has led to the subdivision of the study areainto mapable lithological units. The study area is situatedin the southwestern corner of the Karoo Basin and isbounded by the longitudes 21°19’ and 21°45’ east andlatitudes 32°20’ and 33°00’ south. Initial geological map-ping of the area was accomplished in the laboratory withthe aid of aerial photographs as well as mosaics of GoogleEarth satellite imagery in order to recognize prominentlithological units. This map of the study area was subse-quently checked through fieldwork. Two stratigraphicsections were measured through the entire AbrahamskraalFormation in the study area, and information docu-mented includes lithofacies type, sedimentary structures,grain size, palaeocurrent directions, and ranges of verte-brate fossils. Although no Lower Beaufort lithologicalunits are continuous (Rossouw & De Villiers 1953) in thestudy area, it was possible, based on their lithology, torecognize several prominent stratigraphic units whichextend across the entire study area and have been coded.Biostratigraphically the study area is dominated bydinocephalia and small dicynodonts Robertia, Eosimopsand Diictodon as well as therocephalians. The last occur-rence of dinocephalia in the study area is 21 m below thePoortjie sandstone.

Boonstra, L.D. 1969. The fauna of the Tapinocephalus Zone (Beaufortbeds of the Karoo). Annals of the South African Museum 56, 1–73.

Keyser, A.W. & Smith, R.M.H. 1978. Vertebrate biozonation of theBeaufort Group with special reference to the western Karoo Basin.Annals of the Geological Survey of South Africa 12, 1–36.

Rossouw, P.J. & De Villiers, J. 1953. The geology of the Merweville area,Cape Province. An explanation of Sheet 198 (Merweville). 78 pp.Pretoria, Geological Survey of South Africa.

Investigating the evidence for soft organic tissuepreserved with the MH1 and MH2 Australopithecussediba fossils from the Malapa Cave site

Keeling, Rachelle; Berger, Lee R.Institute for Human Evolution, School of GeoSciences, University of the

Witwatersrand, Johannesburg, South [email protected] / [email protected]

The well-preserved remains of the Australopithecus sedibaspecimens from the Malapa Cave site, in South Africa,might have retained soft tissue in the form of fossilizedskin. A multi-disciplinary approach combining morpho-logical and molecular imaging techniques was utilized toinvestigate whether or not original organics might berecovered. Two primary samples were investigated. Thefirst lay above the posterior surface of MH1’s cranium,while the second was derived from the mandible ofMH2. Six diverse, non-destructive techniques, combiningmicro-CT scans, Raman spectroscopy and more, build aprovocative body of evidence. Any soft original materialdiscovered from the 2 Myr Malapa hominins will be a firstin the field of paleoanthropology.

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Histology of sauropod cervical ribs: implicationsfor neck posture?

Klein, Nicole1; Sander, P. Martin1; Christian, Andreas2

1 Steinmann Institute of Geology, Mineralogy and Palaeontology, University ofBonn, [email protected]

2 Institut für Biologie und Sachunterricht und ihre Didaktik, University ofFlensburg, Germany

The long neck of sauropod dinosaurs is probably themost distinctive feature of their anatomy and one of thekey evolutionary innovations, permitting their uniquegigantism by reducing the energy spent feeding. How-ever, the position in which the neck was held is still con-troversial, with either a nearly horizontal or a distinctlyraised neck. Arguments put forth have centred on theconstraints imposed by the cardiovascular system, themobility of the intervertebral joints, and on biomechanicalconsiderations. Surprisingly, the distinctive and in sometaxa extremely elongated cervical ribs, that can extendover three vertebrae, have not been studied in this regard,although a bracing function has been hypothesized forthe cervical ribs before.

For the first time bone histology of the extremely longcervical ribs of Brachiosaurus/Giraffatitan and Mamenchisaurusas well as the shorter cervical ribs of a diplodocid sauropodis studied in detail. Besides an expected strong remodel-ing by secondary osteons, we found no primary bonetissue in the cervical ribs but there was metaplastic bone.However, most of the microstructure of the anterior andposterior processes of the cervical ribs is made up of longi-tudinal running fibres, identical to those known from themicrostructure of ossified tendons. The longitudinalorientation of the fibres implies that the cervical ribs hadtransported tension forces and clearly contradicts thebracing hypothesis, because in the case of a bracing func-tion of the cervical ribs the fibres would be expected to runvertically. The main function of the cervical ribs as trans-porter for tension forces is also supported by the fact thatthe anterior and posterior processes of the cervical ribsshow morphological structures such as striations andshallow depressions, which indicate multiple and closeoverlap of the long cervical ribs which are also preservedon the fossils. On the basis of our histological results wehypothesize that the processes of the cervical ribs wereonly loosely connected by soft tissue and movable alongtheir long axis against each other; this would make theneck much more flexible than previously thought.

A redescription of a burnetiamorph (Therapsida –Biarmosuchia) from Karoo rocks of Malawi(Poster)

Kruger, Ashley; Rubidge, Bruce; Abdala, FernandoBernard Price Institute for Paleontological Research, University of the

Witwatersrand, Johannesburg, South [email protected] / [email protected] / [email protected]

Exposures of Karoo-age Supergroup are represented inthe northern and southern regions of Malawi. The faunadiscovered in the Chiweta Beds of northern Malawiincludes typical representatives of Late Permian age,which suggest that it is coeval with the CistecephalusAssemblage Zone of the South African Karoo. TheChiweta fauna reported by Jacobs and colleagues in

2005 includes the dicynodont Oudenodon and a newbiarmosuchian. The latter taxon is represented by theskull and lower jaw of a burnetiamorph, a group ofbiarmosuchians that present numerous bosses and swell-ings on the skull. Burnetiamorphs are currently knownfrom seven genera of Middle to Late Permian age, five ofwhich are known form the Beaufort Group of SouthAfrica, the other two from Russian deposits. MAL 290represents the first burnetiamorph from Malawi. Apreliminary description of this material, MAL 290, includ-ing a phylogenetic analysis was presented by Jacobs andcolleagues in 2005, who found the Malawi specimen to bethe sister taxon to Probunetia, Burnetia and Bullacephalus.Additional preparation of the skull of the burnetiamorphfrom Malawi, allows a full description of the material andhas enabled a revised phylogenetic analysis.

Large burrows and palaeoenvironmentalreconstruction of the Early Triassic KatbergFormation, southeastern Main Karoo Basin,South Africa (Poster)

Krummeck, William; Bordy, EmeseDepartment of Geological Sciences, 13 University Avenue, Upper Campus,

University of Cape Town, Rondebosch, 7700 South [email protected] / [email protected]

The Early Triassic Katberg Formation (Lystrosaurus As-semblage Zone) of the Beaufort Group records the periodwhen life recovered from the Permo-Triassic extinctionevent during which nearly 90% of all life was wiped out.Several burrow casts of ~11 cm in diameter and up to 2 min length, and morphologically similar to those describedfrom the Triassic of Antarctica, were found in the KatbergFormation in the Free State and Eastern Cape (SouthAfrica). The latter locality (NW of Cathcart) previouslyproduced several vertebrate fossils and Kitchingnathusuntabeni (holotype BP/1/1187).

This study seeks to: 1) reconstruct the local palaeo-environments, 2) determine the purpose of the burrowsand 3) the possible burrow makers. The analyses of theburrow casts, vertebrate fossils and host sedimentaryrocks have been undertaken through a multidisciplinaryapproach based on ichnological, sedimentological, petro-graphical, stratigraphic, taphonomic and paleontologicalevidence gathered in the field and laboratory observa-tions.

Some of the findings related to the burrows include:diamond-shaped pattern of incised scratches (bioglyphs)on the surface of burrow casts indicating the alternatingscratching directions by the burrow maker; bilobatecross-sectional shape suggesting travel wear due to asemi-sprawling locomotion by the burrow maker; multi-stage passive filling of burrows implying that the burrowswere filled up in several flooding events. Furthermore,a ~0.5 mm calcite rim around some of the casts, the ’-cement-supported’ grain fabric in the burrow fill, thedeformation of burrows by calcite concretions and prefer-ential calcretization of the burrow casts relative to the hostrocks are all indicative of the high permeabilities of theburrow fill before cementation, a process that likelyoccurred within soil profiles in very early diagenesis dueto evaporation. The burrows tend to occur in floodplain

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mudstones overlain by channel fill and crevasse splaysandstones. The close proximity to such sandy depositsand the sedimentary and ichnofacies of the hostmudstones and coarse-grained burrow fills suggest thatthe burrowing occurred in parts of the floodplain thatwere in fairly close proximity to the higher energy fluvialchannels.

With this on-going study, we hope to gain furtherinsights into the earliest Triassic ecosystems of the Karooand the behaviours (e.g. survival strategies) of the organ-isms after ’the Great Dying’ at the Permian-Triassicboundary.

Preliminary analysis of a Triassic dinosaur fromthe Lower Elliot Formation of Lesotho

Krupandan, Emil; Chinsamy-Turan, AnusuyaDepartment of Zoology, University of Cape Town, Rondebosch, 7700 South Africa

[email protected]

The Lower Elliot Formation (LEF) of South Africa is therichest known Triassic assemblage dominated by rela-tively large sauropodomorph dinosaur remains ofAntetonitrus ingenipes, Blikanasaurus cromptoni, Melanoro-saurus readi, Plateosauravus cullingworthi, Eucnemesaurusfortis and Aardonyx celestae. Massospondylus, a prosauro-pod, is most abundant in these deposits, with a number ofindividuals of different ontogenetic sizes (adults, juve-niles, embryos within eggs). We investigate undescribedmaterial from the site, Maphutseng in Lesotho. Duringthe 1950s and 1960s Francois and Paul Ellenberger studiedthe LEF at Maphutseng in the Mohale’s Hoek region andrecovered numbers of presumed prosauropod remainsfrom a bone-bearing lens.

Apart from three cursory descriptions by the Ellenbergersand Gauffre of some of the material, not much work hasbeen done on the bones excavated. Ellenberger &Ellenberger reported that the bones recovered from thesite numbered in the high hundreds and were identifiedas a ’new’ prosauropod similar to a Plateosaurus orMelanosaurus.

We assess the postcranial remains to provide an anatom-ical and taxonomic description. We apply histologicalmethods to ascertain biological aspects of this largesauropodomorph. We hope this multi-dimensionalapproach will provide a better understanding of theMaphutseng dinosaur, and its relationship to contempo-raneous sauropodomorphs in the Lower Triassic of SouthAfrica.

Sereno, P.C. 1999. The Evolution of Dinosaurs. Science 284, 2137–2147.Yates, A.M. 2006. Solving a dinosaurian puzzle: the identity of Aliwaliarex

Galton. Historical Biology, 1–31. 19(1), 93–123Gauffre, F. 1993. Biochronostratigraphy of the lower Elliot Formation

(southern Africa) and preliminary results on the Maphutsengdinosaur (Saurischia: Prosauropoda) from the same formation ofLesotho. In: Lucas, S.G. & Morales, M. (eds), The Nonmarine Triassic.New Mexico Museum of Natural History and Science Bulletin 3, 147–149.

Ellenberger, F., Ellenberger, P. 1956. Le gisement de Dinosauriens deMaphutseng (Basutoland, Afrique du Sud). Comptes Rendus de laSociete Geologique de France 8, 99–101

Reisz, R.R., Scott, D., Sues, H., Evans, D.C., Raath, M.A. 2005. Embryos ofan Early Jurassic prosauropod dinosaur and their evolutionarysignificance. Science 309, 761–764.

Brusatte, S.L., Nesbitt, S.J., Irmis, R.B., Butler, R.J., Benton, M.J., Norell,M.A. 2010. The origin and early radiation of dinosaurs. Earth-ScienceReviews 101, 68–100.

Upchurch, P., Barret, P.M., Galton, P.M. 2007. A phylogenetic analysis ofbasal sauropodomorph relationships: implications for the origin ofsauropod dinosaurs. Special Papers in Palaeontology 77, 57–90.

Insect herbivore diversification after theend-Permian crisis: evidence from leaf miners

Labandeira, Conrad1,2,3; Prevec, Rose2

1 Smithsonian Institution; Washington, DC, [email protected]

2 Rhodes University, Grahamstown, 6140 South [email protected]

3 University of Maryland, College Park, MD 207, U.S.A.

One of the ecological consequences of the end-Permianbiotic turnover was the origin on land of new plantcommunities during the Triassic dominated by cladesoverwhelmingly different from those occurring duringthe Permian. Insects also underwent a major diversifica-tion and new herbivore clades arose, either opportunisti-cally or in tandem with their plant hosts. Nowhere is thisplant-and-insect pattern of radiation and colonizationbetter demonstrated than in the Karoo Basin of SouthAfrica. In an ongoing examination of plant damage infloras spanning an approximately 50-million-year intervalfrom the late Early Permian through the early Late Trias-sic, we have initially selected the colonization patterns ofleaf-mining insects, as evidenced by their distinctive anddiagnosable mines preserved on a broad spectrum ofvascular plant hosts. In the future, we will comparepatterns of plant-host colonization by leaf miners togallers, piercer-and suckers, seed predators, and otherinsect functional feeding groups that exhibit elevatedlevels of host specificity.

Our leaf-mine sub-dataset begins during the latestPermian in which a distinctive blotch mine has beenrecorded on a glossopterid leaf taxon found at a singlelocality in the Emakwezini Formation (NormandienFormation), followed by the absence of leaf mining (or forthat matter, hardly any specialist herbivores) in theAnisian Burgersdorp Formation. However, throughoutthe early Carnian Molteno Formation, seven leaf-miningdamage types (DTs) occur in 30 stratigraphic levels from22 localities. The 27 leaf-mined plant-host species includesphenopsids, ferns, and a particularly rich spectrum ofseed plants comprising broadleaved voltzialean conifers,cycads, five major lineages of ginkgoopsids, corystosperms,and taxa possibly affiliated with bennettitopsids andgnetopsids. Our data preliminarily indicate dietary parti-tioning of foliar tissues, including full-depth and epider-mal mines in leaf blades, rachides and midribs, and rarelyin horsetail stems. Three distinctive mine DTs occur exclu-sively on herbaceous plants, notably horsetails, ferns andkannaskoppiaceous ginkgophytes in Horsetail Marsh andFern Kannaskoppia Meadow habitats. One very abun-dant damage type, DT71, is centred on the broadleavedvoltzialean conifer Heidiphyllum elongatum in HeidiphyllumThicket, but also occurs in other seed-plant lineages,particularly cycads, ginkgophytes and corystosperms,occurring in Dicroidium dominated riparian forests and inDicroidium- and Sphenobaiera-dominated woodland. Thedistinct ecological centre of leaf mining distribution isHeidiphyllum Thicket, from which other habitats were

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possibly colonized. These data indicate an evolutionarilydynamic pattern of plant-host colonization by herbivo-rous insects, paralleling the associational diversity theAngiosperm Revolution that began approximately 100million years later.

The small mammal fauna from Koanaka South,Botswana, and the implications for reconstructingPleistocene environments

Lewis, PatrickDepartment of Biological Sciences, Sam Houston State University,

Huntsville, TX 77340, [email protected]

Recent excavations of fossiliferous deposits from BoneCave have produced a diverse Plio-Pleistocene mammalianfauna. The cave is located in the southernmost section ofKoanaka Hills, northwestern Ngamiland, Botswana, anarea under-represented for the period. A large assem-blage consisting primarily of rodents and shrews wasexcavated. Similar to many of the Plio-Pleistocene smallmammal assemblages in South Africa, taphonomy ofthese fossils is consistent with an owl accumulation andTyto alba (barn owl) is the most likely agent. Our goal wasto better inform on the regional paleoenvironment duringthe Plio-Pleistocene.

Specimens attributed to Mus sp., Gerbilliscus sp., andOtomys sp. were identified, but much of the material couldnot be confidently diagnosed below subfamily based onlyon dentition as has been standard practice in the region; tothe level of genus and, frequently, to species. Comparisonof the Koanaka Hills fossil material with recently trappedmodern specimens from the area and comparative speci-mens from the Ditsong National Museum of Natural His-tory (formerly Transvaal Museum) in Pretoria failed toidentify reliable apomorphies for many clades. This resultsuggests that diagnosing these rodents based on dentitionalone is precarious, particularly for many murine taxa.

In addition, genetic work on the modern Koanakarodents finds genetically distinct cryptic taxa of murinesthat are morphologically indistinguishable. As such, a lackof distinct dental morphologies may not be sufficient torule out multiple taxa. Further morphological analyses,using specimens identified to species by molecularmethods, also suggest that some modern specimens inSouth African museum collections may be misidentified.As these specimens have been the primary source ofcomparative material for the diagnosis of small mammalsin the region, any misidentified specimens in these collec-tions could distort ranges of variation and lead to incor-rect identification of fossil specimens.

Once identifications are made and taxonomic listscreated for a locality the environmental preferences ofmodern taxa are used to guide paleoenvironmentalreconstruction. Misdiagnosis of these taxa obviouslydistorts this reconstruction but, in addition recent exami-nations of modern South African rodent distributionssuggest that many taxa do not appear to be constrained byfactors such as rainfall and temperature. The presumedenvironmental preferences of modern taxa may be poorindicators of past environments, and their use certainly

assumes a level of stasis in preferences that appearsunwarranted.

Our data call into question prior paleoenvironmentalreconstructions in the South African Plio-Pleistocene.Well-defined ranges for dental characters must be estab-lished and clear apomorphies defined for modern SouthAfrican murine taxa relied upon to guide the interpreta-tion of regional fossil specimens. The variables constrain-ing rodent ranges must also be more clearly defined. Untilthen current taxonomic lists and their associatedpaleoenvironmental reconstructions for Plio-Pleistocenelocalities should be used with caution.

Evolutionary developmental model for the origin ofthe turtle shell and a novel functional hypothesisfor the origin of the chelonian lung ventilationmechanism

Lyson, Tyler1*; Bever, Gabe1; Scheyer, Torsten2;Hsiang, Allison1; Gauthier, Jacques1

1 Department of Geology and Geophysics, Yale University, New Haven,CT 06511, [email protected]

2 Palaeontological Institute and Museum, University of Zurich, Zurich, Switzerland

The origin of the turtle shell and the constraints it placeson how turtles breathe are interdependent problems thathave fascinated scientists for the past three centuries. Thediscovery of the stem turtle Odontochelys semitestaceasupports the developmental de novo hypothesis, because itconfirms that the costals and neurals are producedthrough the outgrowth of (sub) dermal bone from theperichondral collar of the dorsal ribs and vertebrae. Thisdiscovery allows for the integration of developmental andfossil data into an evolutionary developmental (evo-devo)model for the shell’s origin that makes explicit predictionsfor the contentious early history of the turtle stem. Weexpand this model by integrating novel anatomical andhistological data for Eunotosaurus africanus, a speciesrecovered as a stem turtle in both global phylogeneticanalyses of amniotes and parareptiles. We consider thephylogenetic signal within a dataset based on shell-relatedcharacters and shelled taxa currently considered as poten-tial relatives of turtles. This analysis tests the hypothesesthat the de novo shell appeared once in amniote historyand that E. africanus is a stem turtle.

Results support the hypotheses that the de novo shellappeared only once in vertebrate evolution and thatE. africanus is a stem turtle. Characters E. africanus shareswith stem turtles include: ten or less elongate dorsal verte-brae, nine pairs of anterior-posterior broadened ribs, lossof intercostal muscles, insertion of muscles on the ventralside of the dorsal ribs, and (sub) dermal outgrowth ofbone from the developing perichondral collar of thedorsal ribs. The successive divergences of E. africanus(broadened ribs; dermal outgrowth of bone from theribs; re-organization of locomotion/respiratory muscles),O. semitestacea (broadened ribs/neurals), and finallyProganochelys quenstedti (fully ossified carapace) results ina sequence of character acquisitions that is fully congruentwith predictions drawn from the evo-devo model.

The evo-devo model further provides a transitionaland mechanistic hypothesis for the origin of the unique

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abdominal muscle lung ventilation system in turtles. Asthe ribs broaden along the turtle stem, they increasinglysupport the abdomen, while simultaneously becomingless effective lung ventilators due to mechanical conflictcreated between adjacent overlapping ribs. As the ribsbroaden, the dual functioning abdominal muscles areslowly freed from their role of providing additionalsupport to the abdominal cavity against torsion and aredelegated into a purely respiratory function. There was ashift from the dual functions found in both the ribs andabdominal muscles of early amniotes, to a division offunction in turtles with the ribs taking on a purely supportrole and the abdominal muscles functioning to only venti-late the lungs.

The avifauna of Langebaanweg (early Pliocene,South Africa)

Manegold, Albrecht1; Louchart, Antoine,2,3

1 Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25,Frankfurt/Main, [email protected]

2 Department of Cenozoic Palaeontology, Iziko South African Museum, Cape Town,South Africa

3 Institut de Génomique Fonctionnelle de Lyon, CNRS, UMR 5242, Université deLyon, École Normale Supérieure de Lyon, 46 Allée d’Italie, 69364 Lyon, [email protected]

The present paper provides an overview on the alreadyfamous, but still insufficiently known avifauna from theearly Pliocene Varswater Formation at Langebaanweg(South Africa). This site yielded one of the largest pre-Pleistocene bird bone accumulations worldwide. Accord-ing to conservative estimates, at least 60 bird species of 25family-taxa are represented here, and most of them markthe earliest record for these taxa on the African continent.Especially diverse is the fossil record of seabirds such aspetrels, prions, and shearwaters (Procellariidae), penguins(Spheniscidae), as well as terrestrial taxa such as shore-birds (Charadriiformes) and songbirds (Oscines,Passeriformes). Recent identifications of new bird specieshave shed new light on current hypotheses on thepalaeoenvironment and palaeoecology at Langebaanweg.For instance, honeyguides and several cavity nesting birdssuch as parrots and woodpeckers indicate the presence offorested habitats, but evidence for two species ofsandgrouse (Pteroclidae), a taxon generally associatedwith arid to semi-arid landscapes, reveal a little knowncomponent of the palaeoenvironment of Langebaanwegarea during the Early Pliocene.

Bone microstructure of metapodial bonesreveals growth pattern and life history featuresof Miocene Hipparion from Spain

Martinez-Maza, Cayetana1*; Alberdi, Maria Teresa1;Gomez, Santiago2; Prado, Jose Luis3

1 Department of Palaeobiology, Museo Nacional Ciencias Naturales (CSIC),Madrid, [email protected]

2 Department of Pathological Anatomy, University of Cádiz, Cádiz, Spain3 INCUAPA, Departamento de Arqueología, Universidad Nacional del Centro,

Del Valle, Olavarría, Argentina

Hipparions characterizes the Upper Miocene and Plio-cene faunas of Eurasia and Africa. They represent anintermediate stage in horse evolution towards highercrowns, larger size and reduced autopodials. They present

an autopodial composed by a major central toe and tworeduced lateral ones. Their morphology has been studied,but until now not histologically. Bone microstructurereflects development, growth and adaptative changesduring life. These aspects of the life history of an extinctanimal, can be assessed as fossil bone generally maintainsits histological integrity. We characterize the micro-structure from 50 central and lateral metacarpal and meta-tarsal bones of Hipparion concudense from the Vallesian andTurolian fossil sites of Valles de Fuentidueña (Segovia)and Concud (Teruel) of Spain. All analysed bones havefused epiphyses. Bright field and circular polarized lightmicroscopy of transverse ground sections of midshaftsreveals that central metapodials are formed of corticalcompact bone made of primary bone tissue with variableextension of secondary bone. Primary bone tissue is com-posed by many layers of primary osteons formed duringgrowth by periosteal apposition. Secondary bone showsseveral generations of haversian osteons which mainlydisplay a hooked and alternate collagen pattern; oneindividuals show multiple primary osteon system andfew secondary osteons in the posterior area. Other speci-mens show similar histology but haversian bone extendsboth in the posterior and the inner area of the dorsalregion. Some specimens show secondary osteonsthroughout the cortical bone. The microstructure ofleteral metapodials is similar to that in the centralmetapodials, but in the lateral one the marrow cavity isreduced to a small space or completely absent. Primaryand secondary bone is present in variable proportions. Inparticular, large areas of secondary bone are found in themedial side facing the central metapodial. Lines ofarrested growth (LAG) in primary bone in the corticalbone of central and lateral metapodials, may have formedon an annual basis. The number of LAGs is variable; in theless remodeled cortical bone of a few young individualswe identified from 4 to 6. Structural changes in the corticalbone suggest different age-related bone maturation asso-ciated with broad developmental stages in Hipparionconcudense. At early stages of development, metapodialbones show appositional periosteal associated with fastgrowth together with secondary osteons in the posteriorregion. In young specimens, the short distance betweenLAGs and the increase of haversian bone area couldindicate a decelerating growth. The adult specimenspresent completely remodeled cortical bone. Secondaryosteons in the central metapodials of young individualsoccur in areas associated with lateral metapodials. Lateralmetapodials may inter-relate biomechanically with thecentral metapodials.

Using geometric morphometrics to clarify therelationship between modern soricid and muridtaxa and fossil species from Langebaanweg (5.2 Ma)

Matthews, Thalassa1; Stynder, Deano D.2

1 Iziko South African Museum, Cape Town South [email protected]

2 Department of Archaeology, Faculty of Science, University of Cape Town,Rondebosch, South [email protected]

We used geometric morphometrics to compare mandib-

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ular shape and size of two extinct forest shrew (Soricidae)species found at Langebaanweg (LBW) with extantspecies, to elucidate the relationship between the upperfirst molars from rats (Muridae) belonging to the genusAethomys, which were represented by an anomalouslylarge number of species and/or morphs.

Soricids: This research represented the first study of thesoricid community from LBW, which is represented byfour species. Using geometric morphometrics, the ascend-ing rami of the two Myosorex fossil species were comparedwith each other, and with the extant western and easternCape species Myosorex varius and M. cafer, as well as theelusive M. longicaudatus, which was discovered only in thelate 1970s in the Knysna area, south coast. The mandiblesof the fossil species are characterized by ascending ramiwhich are relatively gracile and differ in shape and sizefrom modern species, and the existence of a number ofprimitive mandibular and dental features. This evidencewas interpreted as indicating that the LangebaanwegForest shrews represent an archaic lineage now extinct insouthern Africa.

Murids: Initial work described two Aethomys species(Aethomys adamanticola and Aethomys modernis) at LBW.Subsequent research led to the secure identification of athird, as yet, undescribed species, and in addition, threemorphs of existing species were also recognized whichshowed sufficient differences to raise the possibility thatthey were different species. Geometric morphometricswas used to explore the relationship of fossil species andmorphs with extant Aethomys species, and to assess theintra and inter-specific variation in the size and shape ofmodern and fossil Aethomys specimens. Geometricmorphometrics indicated a marked similarity in the upperfirst molar (M1) shape between all the analysed species,extant and extinct, and suggests the persistence, over asignificant period of time, of a prototype Aethomys M1shape. The relative warp analyses indicated someinterspecific variation between the two modern species,Aethomys chrysophilus and Aethomys namaquensis, andsome overlap in shape, although they differed signifi-cantly from one another in terms of size. Variability interms of size and to a minor extent, shape, was alsoevident in the fossil Aethomys, and it was concludedthat the various fossil morphs investigated representedintra-specific variability in size and shape rather thanintra-specific sexual dimorphism, or a new Aethomysspecies.

Assessing an exhibition and hands-on fossilfuel workshop as a tool to promote learners’understanding of palaeontology

McKay, IanSchool of Geosciences, University of the Witwatersrand, Johannesburg,

South [email protected]

Public understanding of the possible role of humanactivities in causing climate change and the need for sus-tainable energy sources to mitigate this change is relatedto their understanding of the origin, use of, supply andhazards associated with fuels. However, a proper under-standing of fossil fuels requires a basic knowledge of Earth

history, geological time, evolution, and chemistry whichis taught at schools. In South Africa palaeontology, climatechange and fossil fuels are covered at both primary andsecondary levels. This paper describes the efficacy of aprogramme on fossil fuels presented to primary and highschool pupils at a Science Centre in South Africa as part ofa Sustainable Energy Week. The programme consisted ofa fossil fuel exhibition with a five minute overview onfossil fuels presented by a postgraduate student. In addi-tion there was a hands-on workshop consisting of a pre-sentation explaining the origin of fossil fuels, futuresupplies, and their probable role in influencing climatechange. Students compared crude oil with engine oil,as well as peat with different types of coal and alsoused polystyrene balls to model hydrocarbons. Pre- andpost-workshop questionnaires were used to assessthe programme. Results suggest that the learners’understanding of fossil fuels and their possible relation-ship to climate change is poor, resulting in their opin-ions being shaped by societal pressure rather thanactual understanding of the concepts involved. Theprogramme was partially successful in assisting learnersto understand fossil fuels and their relationship to climatechange.

The anatomy and phylogenetic relationshipsof Antetonitrus ingenipes (Dinosauria,Sauropodomorpha) from the Late Triassicof South Africa: implications for the originsof Sauropoda

McPhee, BlairBPI Palaeontology, University of the Witwatersrand, Private Bag 3, WITS 2050,

Johannesburg, South [email protected]

The sauropods were the dominant herbivorous dino-saurs for the majority of the Mesozoic, representing thelargest terrestrial animals to have ever evolved. How thegroup rose to such an important role is a question of muchinterest. However, until recently the anatomy – includingthe basic osteological framework of the most basalmembers of the Sauropoda had remained something of amystery. Exacerbating the problem was a scarcity of goodspecimens and a lack of consensus on the wider relation-ships of the Sauropoda among the more inclusivesauropodomorph clade. The situation has improved inrecent years, and new discoveries from Thailand, Argen-tina, China, and South Africa have begun to providepalaeontologists with the means of unravelling the originsof this fascinating group of animals. One of these discov-eries, Antetonitrus ingenipes from the lower Elliot Forma-tion of South Africa, preserves significant elements of boththe axial and appendicular skeleton and therefore repre-sents one of the most significant early sauropoddiscoveries. Dating to the Norian of the late Triassic,Antetonitrus displays an intermediate morphologybetween later sauropods and their `prosauropod’ fore-bears. This intermediate morphology and early ageuniquely informs us on the pattern and timing of theacquisition of numerous sauropod traits, and severalrecent phylogenies have resolved Antetonitrus as the mostbasal of all Sauropoda. However, our understanding of

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basal sauropodomorph interrelationships – particularly atthe sauropodomorph/sauropod transition – continues toremain controversial, and the full informational signifi-cance of Antetonitrus is yet to be fully explored. A compre-hensive description of the Antetonitrus fossil assemblage istherefore considered a necessary and timely undertakingin light of the current state of basal sauropodomorphstudies, and will form the essence of my current Mastersof Science project at the University of the Witwatersrand.Additionally, as the character states of Antetonitrus haveyet to be based upon a full osteological description, arevised cladistic analysis may yield additional insight intothe precise evolutionary context of this important transi-tional taxon, especially in light of the recent advances insauropodomorph phylogenetics.

Note on the fossil fauna and flora at OngongoSprings, Damaraland(Poster)

Mocke, HelkeNational Earth Science Museum, Geological Survey of Namibia,

Windhoek, [email protected]

Knowledge of Quaternary-aged fossils in Namibia issomewhat lacking, although several such sites were iden-tified, for example, by Hermann Korn and Henno Martin.The community-run Ongongo Springs in WesternNamibia yielded thick layers of tufa (freshwater carbonate)containing a rich collection of impressions of macroscopicfossil plant leaves, roots, branches and trunks. The leafimpressions show mostly primary venation, making theiridentification very difficult. The absence of organic materialwithin the preserved leaves, roots, stems and trunksmakes C14 dating impossible. However, dating of the tufaswill ascertain when the plant matter was covered by theprecipitated material. A possible sedge leaf was noted andimpressions of what appear to be those of the sycamorefig, Ficus sycomorus (Family Moraceae) and Mopane,Colophospermum mopane (Family Fabaceae) leaves. How-ever, no fruits or seeds were found. To date only a singlevertebrate fossil has been reported by a visitor to thesprings. It has been suggested that it is the impression of afrog skeleton. Land snails were observed in the tufas andsurrounding calcretes and are comparable to the modernspecies Sculptaria in general morphology. No dating has asyet been done on the tufa, due to an absence of datingequipment in Namibia.

Microstructural differences in femoral bone ofAbrothrix longipilis (Cricetidae, Sigmodontidae)populations from different altitudes(Poster)

Montoya, Germán1; Iturra-Cid, Myriam1; D’Elía, Guillermo2

1 Facultad de Ciencias Naturales y Oceanográficas, Universidad de Concepción,Concepció[email protected] / [email protected]

2 Instituto de Ciencias Ambientales y Evolutivas, Universidad Austral de Chile,Casilla 567, [email protected]

Bone is a plastic and highly responsive tissue thatchanges in response to different mechanical and environ-mental conditions. Considering this, bone microstructure(BM) of vertebrates is useful to assess the ecological and

life history characteristics of extant and extinct species.However, accurate knowledge about geographic andclimatic variation on the arrangement of cellular andstructural components of bone is scarce; we do not knowthe effects of high-elevation conditions e.g. hipoxy andderivated process as angiogenesis and osteoclastogenesis,upon BM. Knowing these effects on BM could be very use-ful to identify and estimate ecological and paleoecologicaltraits in extinct assemblages (bonebeds) associated withorogenic uplifts. We compare BM from populations exposedto different environmental pressures associated with at-mospheric oxygen availability at low and high elevations.

We assessed quantitative and qualitative differences inbone vascularization in populations of long-haired grassmouse Abrothrix longipilis from two localities; one atmoderate elevation (n = 5; 1420 m) and another at sealevel (n = 5; 15–32 m). We hypothesized that the hypoxicconditions encountered by the high elevation population(He) would be reflected by the presence of moreHaversian systems and vascular canals than the sea-levelpopulation (Le).

Osteohistological sections of f emora were made, andprepared. BM was described in terms of the number ofHaversian systems and vascular canals structures, inaddition to description of their arrangement and pattern.A nonparametric Mann-Whitney U-test was used to com-pare whether the mean values corresponding to the Heand Le samples were significantly different (P <0.05).

Populations differed in the proportion and frequency ofthe microstructural components and in the arrangementand distribution of such elements. Similar to other rodentsand mammals, both populations exhibited a small num-bers of Haversian systems (38) and a considerable abun-dance of vascular canals (242). Individuals of Le (5.8 ± 4.14,P = 0.028) had a tendency to have more Haversian sys-tems than those of He (1.8 ± 1.48). Inversely, He individu-als have a higher frequency of vascular canals (39.6 ±19.91, P = 0.036) than Le (8.8 ± 3.27). It is likely that theincreased frequency of vascular canals is associatedwith increased blood supply in populations at highand moderate altitude, where oxygen levels are lowerthan at sea level. Moreover, the slightly larger number ofHaversian systems in the Le opposed to the He populationindicates possible differential metabolic dynamics in boneremodeling associated with altitudinal gradients.

There are significant differences in the BM pattern ofA. longipilis at different altitudes and BM could be used asa proxy for high altitude environments. This is an interest-ing and useful tool for interpreting adaptive and paleo-ecological aspects of organisms related to topographicvariables such as in the Andean Altiplane.

The hominin thoracic shape

Nalla, Shahed1,2,3; Zipfel, Bernhard3

1 Department of Human Anatomy and Physiology, Faculty of Health Sciences,University of [email protected]

2 Institute for Human Evolution, University of the Witwatersrand3 Bernard Price Institute for Palaeontological Research, University of the

Witwatersrand, Johannesburg, South [email protected]

Understanding the locomotory and postural changes

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that affected the early hominins is important in order togain an understanding of primate evolutionary processes.It has been hypothesized that, specifically amonghominins, major evolutionary changes have taken placein rib form and structure, which, in turn influencedthoracic shape. Examination of the partial thorax of theAustralopithecus afarensis skeleton (’Lucy’) had contra-dicted the previous suggestion that the thoracic shape ofthis species was more ’ape-like than human-like’. Thisprompted significant debate and highlighted the impor-tance of understanding and interpreting thoracic shape inthe fossil hominin record. Earlier ideas of the dimensionand shape of the hominin thorax and its evolution havebeen further challenged by subsequent studies of themore-complete thoracic remains of the Kebara 2 Neander-thal and the Nariokotome Homo erectus juvenile.

The complex geometric and dynamic relationshipbetween the thorax, the clavicle and the scapula has alsonever been fully understood, even in humans, let alone itsvariation and implications among primate species withdiverse locomotor behaviours.

The present study undertook to determine the shape ofthe hominoid thorax and its influence on the position ofthe upper limb. The morphology of the thoracic and theupper limb elements (the proximal end of vertebro-sternal ribs, the sternum and associated costal cartilages,the clavicles and the scapulae) of selected extant primatespecies was analysed by taking 21 linear measurements(from which 12 indices were derived so as to quantifymorphometric information from homologous points).These measures were then compared to those found inthe fossil record in order to determine whether there areany structural correlates between the extant and theextinct hominin species.

Preliminary results indicate a generalized consensusthat modern humans exhibit a narrow upper and lowerthoracic shape, when compared to the primitive ape-likethorax which is narrow at the top and wider at the bottom.The observed thoracic shapes facilitate optimal position-ing of the upper limb for efficient locomotion and behav-ioural requirements.

The coprolites from the Early Triassic fossil siteDriefontein, eastern Free State

Neumann, Frank H.1,2; Yates, Adam2,3; Hancox, John2

1 Forschungsstelle Palaeobotanik, University of Münster, [email protected]

2 Bernard Price Institute for Palaeontological Research, University of theWitwatersrand, Johannesburg, South [email protected]

3 Museum of Central Australia, Alice Springs, [email protected]

The Driefontein locality comprises lacustrine muds andfluviatile sandstones of the Burgersdorp Formation,which was deposited in the main Karoo Basin from theEarly Triassic through to the early Middle Triassic. The fos-sil site is of special importance since it shows one of theworldwide rare examples of Early Triassic recovery faunafollowing the End-Permian extinction event. These harshconditions are reflected by the depauperate nature of

Early Triassic ecosystems and the small sizes of theirconstituent species, a phenomenon known as the Lilliputeffect.

Our aim is to contribute to a better understanding of thepalaeoecology by looking at the coprolites in a thinbonebed, which are 400 µm to several centimetres indiameter with an array of methods including lightand electron miscroscopy, thin section and silicon peels.Hybodont sharks, lungfish, several temnospondyl amphibi-ans and a large basal archosauriform, all evidenced at thesite by fossil findings, are possible producers of the copro-lites.

We documented numerous fossils which were withinthe coprolites, including an insect wing impression,dipnoan tooth plates, arthropod remains, bone frag-ments, fish scales, dermal bones and plant remains.Among the most outstanding findings are bivalve impres-sions which might belong to unionids although they areunusually small and the identification is hampered bylimited preservation. If the specimens do representunionoids then these Early Triassic examples may be anexample of the Lilliput effect. Since the oldest incontro-vertible unionoids are also from sub-Saharan Africa, it ispossible that this subcontinent hosted the initial radiationof the Unionoida.

Our findings demonstrate the importance of coprolitesas microenvironments of exceptional preservation thatcontain fossils of organisms that would otherwise haveleft no trace.

A proposed palynostratigraphic scheme for theKaroo Supergroup of South Africa

Nicoletti, Natasha; Bamford, Marion; Rubidge, BruceBernard Price Institute for Palaeontological Research, University of the

Witwatersrand, Johannesburg, South [email protected]

The Karoo Basin is an excellent source of palaeobotanicalfossils, including palynomorphs, but previous researchhas concluded that the abundance of palynomorphs insome Karoo rocks is not sufficient to establish a bio-zonation for the entire Karoo sequence. Recent extensivesampling from the Karoo stratigraphic succession hasyielded numerous palynomorphs, and it appears possibleto generate a palynostratigraphic scheme for the Karoosedimentary sequence from both the proximal and distalfacies (Catuneanu et al. 1998). The palynological signa-tures of the Ecca Group, Permian Beaufort Group, TriassicBeaufort Group, Molteno Formation and Elliot Formationare described. The recent determination of radiometricdates for most of the Permian vertebrate biozones of theBeaufort Group (Rubidge et al. 2010) will enable precisedating for some palynological zones. Comparison of theKaroo palynological biozonation with the existing Austra-lian scheme and other Gondwana palynomorphs revealssimilarities at the generic level but South Africa possessessome unique species that have not been recovered fromother Gondwana deposits. The taphonomy of palyno-morphs may prove useful for palaeoenvironmental deter-mination within different deposits of the Karoo Supergroup

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and is expected to corroborate geological evidence ofPermian and Triassic palaeo-environments.

Catuneanu, O., Hancox, P.J. & Rubidge, B.S. 1998. Reciprocal flexuralbehaviour and contrasting stratigraphies: a new basin developmentmodel for the Karoo retroarc foreland system, South Africa. BasinResearch 10, 417–439.

Rubidge, B.S., Erwin, D.H., Ramezani, J., Bowring, S.A. & de Klerk, W.J.2010. The first radiometric dates for the Beaufort Group, KarooSupergroup of South Africa. Proceedings of the 16th Conference of thePalaeontological Society of southern Africa, Howick, 5–8 August 2010.

Variation of cranial morphology and size in thegorgonopsian Aelurognathus

Norton, Luke; Rubidge, Bruce; Abdala, FernandoBernard Price Institute for Palaeontological Research, Palaeosciences Centre,

University of the Witwatersrand, Johannesburg, South [email protected]

The Gorgonopsia were a group of highly specializedcarnivores that lived approximately 265–250 Ma ago,during the Late Permian. Skulls belonging to Gorgo-nopsia range in size from those of a domestic cat, to largerthan any extant terrestrial predator. Despite this widerange in size, morphological variat ion of thegorgonopsian skull is relatively conservative. The aim ofthis study was to identify and describe the extent of mor-phological variation occurring in a sample of gorgo-nopsian skulls attributed to the genus Aelurognathus.Aelurognathus represents the gorgonopsian genus with 16assigned specimens which are attributed to six species.The length of the skull in this sample varies from approxi-mately 18 cm to 34 cm. Specimens were examined in orderto elucidate intra and interspecific variation. Differencesobserved at the interspecific level allowed for the recogni-tion of three morphotaxa, based predominantly on char-acters of the skull roof. These characters included the stateof a preparietal, and a contribution of the frontal to thesupraorbital margin. A high degree of variability in themorphology and state of the preparietal has been welldocumented in various taxa of extinct and extant synapsids.Sutural boundaries between elements of the skull roofhave also been demonstrated to be highly plastic. Theextent of sutural variation has been shown to differ con-siderably, even between opposite sides of the same skullin a specimen of lemur. Taking these observations intoaccount, the hypothesis that all specimens represented asingle taxon exhibiting a high degree of morphologicalvariation was tested using allometric techniques.Twenty-seven linear measurements were chosen, suchthat variability in skull size and shape could be accountedfor in all dimensions. Results of the bivariate allometricanalyses showed a high level of correlation of the plottedmeasurements with the calculated best-fit lines, support-ing the single taxon hypothesis. While Aelurognathus haspreviously been divided into six species using only mor-phological characters, this study has shown that the char-acters used exhibit a high amount of variability. Excessivetaxonomic importance has been placed on characters thatnow seem trivial, leading to a high number of describedspecies of Aelurognathus. As such, it is proposed that allspecies currently attributed to the genus Aelurognathus besynonymized with the type, Aelurognathus tigriceps.

Establishing incidences of dental calculus andassociated plant microfossils in South AfricanPlio-Pleistocene hominin dentition

Odes, Edward1; Berger, Lee2; Henry, Amanda3;Bamford, Marion1

1 Bernard Price Institute for Palaeontological Research, School of Geosciences,University of the Witwatersrand, Johannesburg, South [email protected] / [email protected]

2 Institute of Human Evolution, School of Geosciences, University of theWitwatersrand, Private Bag 3, Johannesburg, 2050 South [email protected]

3 Max Planck Institute for Evolutionary Anthropology, Leipzig, [email protected]

Study of the dental remains from Malapa, a fossil-bearing karstic cave site located in the Cradle of Human-kind (Berger et al. 2010), have revealed the presence ofdental calculus and associated phytoliths preserved onthe teeth of Australopithecus sediba (MH1) (Henry et al.,in press). This raises the possibility that dental calculusand plant microremains may be present on homininmaterial from other cave sites, where fossils are preservedunder similar conditions. Our aim was to establishwhether dental calculus is present on the teeth of othersouthern African Plio-Pleistocene early hominins, and ifso, how widespread it is, and how it manifests. We exam-ined the dental collections of the Plio-Pleistocene agegroup of hominin sites of South Africa. For those fossils onwhich we found adherent material, we performed severalanalyses to determine whether this material is calculus ornot.

As in previous studies, we performed a visual examina-tion of the location and appearance of the calculus, andwhere possible, a comparison with the texture of the sedi-ment matrix surrounding the fossil. In addition, weemployed other methods, such as scanning electronmicroscopy with energy dispersive X-ray spectroscopy(SEM/EDX), which can establish whether the material waschiefly calcium phosphate (calculus) or something else. Incertain cases, we removed small quantities of this materialto determine whether it included food particulates andmicrofossils. In these cases, we also looked for microfossilsin the surrounding matrix as a control. For those fossils onwhich no calculus material was found, we explored thepossible causes for the lack of calculus, including preser-vation environments and potential preparation.

The results from this study will benefit future analysis byproviding a list of target fossils, and a protocol for estab-lishing the presence of calculus. These future studies maybe able to obtain direct evidence of consumed food thatcan directly be associated with the individual hominins’feeding behaviours. This could result in significant cluesto the diet and ecology of not only individual hominins,but populations, species and comparisons of diet andbehaviour between species and genera.

Berger, L.R., de Ruiter, D.J., Churchill, S.E., Schmid, P., Carlson, K.J.,Dirks, P.H.G.M. & Kibii, J.M. 2010. Australopithecus sediba: a new speciesof Homo-like australopith from South Africa. Science 328, 195–204.

Henry, A.G., Ungar, P.S., Passey, B.H., Sponheimer, M., Rossouw, L.,Bamford, M., Sandberg, P., de Ruiter, D.J., Berger, L. 2012. The diet ofAustralopithecus sediba. Nature 487, 80–93.

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Application of spore-pollen biostratigraphy in thedeep offshore Cenozoic Niger Delta

Olayiwola, Moshood; Bamford, MarionBernard Price Institute for Palaeontological Research, Palaeosciences

Centre, School of Geosciences, University of the Witwatersrand,Johannesburg, South [email protected] / [email protected]

The application of spore and pollen biostratigraphy inthe deep offshore Miocene to Pliocene Niger Delta strataare of immense importance especially when other taxamake little or no significant impact. Much research hasbeen done on the onshore deposits but the offshoredeposits are now the focal point as these oil and gasreserves are now being exploited.

The purpose of this study is to analyse ditch-cutting off-shore samples for their palynomorph contents to create apalynomorph biostratigraphic framework to complementthe existing microfossil and nannofossil biostratigraphy inthe Niger Delta. The published and unpublished litera-ture on the palynology of Cenozoic sediments has beenreviewed and synthesized with a view to identify anddescribe all the offshore palynomorphs in order to estab-lish pollen biozones and ages of strata penetrated by thestudied well A.

Biostratigraphic studies of spores and pollen recentlycarried out on well A from the southeastern Niger deltabasin resulted in identification of over 112 spores andpollen taxa with their respective frequency distributions.This in turn resulted in a palynostratigraphic zonation ofthe strata penetrated by well A. Three distinctive zonescomprise of seven subzones, ranging from Upper Mio-cene to Lower–Upper Pliocene Series, were established inthis study.

The Upper Cretaceous calcareous nannoplanktonof the St Lucia Formation at Nibela Peninsula,KwaZulu-Natal(Poster)

Ovechkina, Maria1,2,3; Mostovski, Mike2,3,4

1 Discipline of Geology, School of Agricultural, Earth and Environmental Sciences,University of KwaZulu-Natal, Durban, South [email protected]

2 Paleontological Institute, Russian Academy of Sciences, Moscow, Russia3 KwaZulu-Natal Museum, Private Bag 9070, Pietermaritzburg, South Africa.

[email protected] School of Life Sciences, University of KwaZulu-Natal, Scottsville, South Africa

The Upper Cretaceous St Lucia Formation at NibelaPeninsula is represented by poorly cemented silty finelygrained sandstones with distinct layers of concretions.The beds are rich in invertebrate fossils and have yieldedvarious cephalopods, bivalves, gastropods, scaphopods,rare echinoids and brachiopods.

One of the sections (S27°59.555’ E32°25.533’) waslithologically described and sampled for calcareousnannoplankton at 0.3–0.5 m intervals. Nineteen bedswere recognized and 40 sediment samples were collected.Subsamples (ca. 100–200 g) were crushed, and smearslides were made for further identification. Calcareousnanno-fossils were identified using a Zeiss Axioskop mi-croscope with crossed nicols at ×1200–1920 magnifica-tion. Almost the entire section contains a rich fossilassemblage, including 80 species with moderate to poor

preservation of nannoplankton. The most common spe-cies are Watznaueria barnesae, Gartnerago obliquum,Reinhardtites anthophorus, Micula decussata, Eiffellithusturriseiffelii , Tranolithus orionatus , T. manifestus ,Lithraphidites carniolensis, Microrhabdulus decoratus, M.belgicus, Prediscosphaera grandis, and Cretarhabduscrenulatus. Less common are Eiffellithus eximius,Cribrosphaerella ehrenbergii, Broinsonia matalosa, Calculitesobscurus, Lucianorhabdus cayeuxii, Biscutum magnum, andArkhangelskiella cymbiformis, whereas Broinsonia parcaparca, B. p. constricta, B. p. expansa, Marthasterites furcatus,and Orastrum campanensis are rare.

According to Perch-Nielsen (1985), the lowermost part ofthe section (Beds 1–3) is referred to the transitional UpperSantonian–Lower Campanian Zone CC17 due to the pres-ence of Calculites obscurus, the first occurrence characteriz-ing the lower boundary.

The Lower Campanian Zone CC18 from bed 4 to themid. bed 15. Its lower boundary is identified by the first oc-currence of Broinsonia parca parca and its upper boundaryis defined by the last occurrence of Marthasterites furcatus.Bed 4 and the lower part of bed 5 belong to SubzoneCC18a due to the presence of Broinsonia parca parca, thefirst occurrence of which is in sample 4. The interval fromthe upper part of bed 5 to the middle part of bed 15 be-longs to undivided Lower Campanian Subzones CC18b–cwith Broinsonia parca constricta, the first occurrence ofwhich is in sample 7. Ceralithoides verbeekii, which occur-rence defines the lower boundary of Subzone CC18c, isnot found.

The upper section, from upper Bed 15 to Bed 19 is theLower Campanian Zone CC19, the lower boundary beingidentified by the disappearance of Marthasterites furcatus.

The calcareous nannoplankton offers an older age of thesediments than it has been suggested previously byammonites and inoceramids.

The usual suspect: experimental taphonomyvindicates the humble dermestid

Parkinson, Alexander HaigBernard Price Institute for Palaeontological Research, University of the

Witwatersrand, Johannesburg, South [email protected]

The family Dermestidae of the order Coleoptera and thegenus Dermestes, was first described by Linnaeus in 1758.The involvement of Dermestes in the animal decomposi-tion process typifies members of this genus. The oldest re-corded occurrence of Dermestidae dates back to the LateTriassic, but they are more reliably found in Cretaceousambers. Dermestes ater, Dermestes carnivorus, Dermestesfrischii and Dermestes maculatus are known to modify bone,but not substantially. Despite this known trait little workhas been done to record the distinctive ways in whichDermestes modify bone. As a result a large body ofpalaeontological literature has arisen in which they areinferred as agents of bone modification without actualisticdata to support such claims. This prompted our investiga-tion in which bone specimens of various types, conditionsand levels of preservation were exposed to Dermestesmaculatus. Experimental tanks were established in a dedi-

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cated insectary which maintained a constant temperatureof 28°C at 40% humidity. The bone specimens were ex-posed to 100 larvae and 35 adults for a period of fourmonths. Specimens were subsequently removed and ana-lysed using an Olympus SZX 16 Multifocus microscopefitted with a digital camera at magnifications between ×7and ×115. The modifications identified in this studywhere then classified, described and compared with thoseattributed to dermestids in the palaeontological literature.We show that the distinctive ways in which bones aremodified by dermestids (bone destruction, irregular sur-face tunnels on cortical bone, pits without associatedstriations on cancellous bone, surface pits with associatedstriations on cortical bone as well as distinctive gnawing-related damage in the form of clusters of multiplestriations) as well as the distribution thereof (periostealsurfaces, along edges, on cortical or only present oncancellous bone, etc.), are not comparable to modifica-tions ascribed to dermestids in the palaeontological litera-ture. Hence, experimental taphonomy has vindicated thehumble dermestid as the modifier of many fossil remainsin prehistory. However, the question still remains, who isresponsible for the modifications described in existingliterature? Ongoing actualistic research, based on theprinciple of uniformitarianism, promises to identify theagent(s) in question, but one has to consider thatdermestid behaviour may have changed, or that the re-sponsible agent is extinct.

Anisian (Middle Triassic) archosaur assemblagesfrom Antarctica, Tanzania and Zambia: how dothey relate to the Karoo?

Peecook, Brandon1; Sidor, Christian1; Nesbitt, Sterling2;Angielczyk, Kenneth3; Smith, Roger4

1 Burke Museum and Department of Biology, University of Washington,Seattle, [email protected]

2 Department of Biology, University of Washington, Seattle, U.S.A.3 Field Museum of Natural History, Chicago, Illinois, U.S.A.4 Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa

The beginning of the Triassic (~252–237 Ma) was a criti-cal interval in vertebrate history. Our understanding ofhow terrestrial ecosystems across southern Pangaearecovered from the end-Permian extinction is basedalmost exclusively on the well-sampled Karoo Basin ofSouth Africa. Southern Pangaea has historically beenviewed as relatively homogenous in community compo-sition, especially during the Late Permian and EarlyTriassic, because sampling outside the Karoo yielded thesame or similar taxa. However, paleontologists have quali-tatively noted that by the Middle Triassic new clades hadradiated and dissimilarity of taxonomic assemblagesacross the supercontinent had become a major feature ofterrestrial ecosystems.

We sampled Middle Triassic localities in the NtawereFormation of Zambia, Manda Beds of Tanzania, andFremouw Formation of the Transantarctic Mountains andhave assemblages that greatly differ from those of theKaroo Cynognathus Assemblage Zone, subzones B and C.Perhaps most significantly, these assemblages containmembers of crown-group Archosauria, which is not

recovered in the better sampled Karoo. From Zambia wehave teeth, a large vertebra belonging to a paracroco-dylomorph pseudosuchian, a proximal femur of the firstshuvosaurid poposauroid outside of the Late Triassic ofthe Americas, and postcranial material of a new genus ofsilesaurid dinosauriform. From Tanzania there is materialfrom a number of pseudosuchians including representa-tives of aetosaur and ctenosauriscid lineages, as well as asilesaurid dinosauriform, Asilisaurus kongwe. FromAntarctica we have a bizarre pseudosuchian femur of thefirst crown archosaur from the Triassic of the continent.These records support the idea that far from being homog-enous, Pangaea was regionally diverse and supportedprovincial assemblages.

We have developed new metrics in biogeographicanalysis to quantify assemblage dissimilarity, focusing onco-occurrences of taxa across basins and applying net-work methods. We have mapped terrestrial assemblagesacross distinct regional bins in southern Pangaea through-out the Triassic and can track increasing assemblagedissimilarity via the increased modularity score in ourtaxonomic co-occurrences. Modules within a network canbe assigned to meaningful groupings in the real world(e.g. geographic location). Traditional biogeographicmethods, including Simpson’s Coefficient, Bray-Curtisindex, and other distance-matrix methods support thecontention that the Karoo Basin is not representative ofthe composition of taxonomic assemblages across southernPangaea during the Anisian. To better understand theradiation of important Mesozoic clades, like Archosauria,we must look to traditionally under-sampled areas.

New small skeletal fossils from the Terrenuvian(Early Cambrian)-aged Sellick Hill Formation(Normanville Group) of South Australia

Penn-Clarke, Cameron; Yates, AdamBernard Price Institute for Palaeontological Research, University of the

Witwatersrand, Johannesburg, South [email protected]

The Cambrian Period is a critical time in Earth’s historywhere fossils of all major phyla of the Metazoa made theirfirst tangible appearance in the rock record. A major factorfor this phenomenon is tied in with advent of bio-mineralization, with the development of harder chitinousor calcareous or phosphatic exoskeletons, i.e. a scleritomecomposed of many sclerites, or shells over their softbodies, and, as seen in later vertebrates, a cartilaginous orphosphatic endoskeleton, possibly as a line of defence.Individual skeletal elements usually are disarticulatedfrom their original position in life, making anatomicalinterpretations and phylogenetic relationships of thesemetazoans obscure. Collectively, these skeletal elementsare referred to as small skeletal fossils (SSF). Estimatingthe phylogenetic relationships of these SSF’s is plaguedby discrepancies in calibrating early Cambrian time.Formal global epoch and stage subdivision schemes forthe Cambrian Period have been suggested, but are poorlydefined and not yet widely implemented. Sedimentarystrata of the eastern Officer and Warburton Basins as wellas the Stansbury and Arrowie Basins and Stuart andSpencer Shelves of the uppermost Adelaide Rift Complex

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of South Australia are well known to contain fossils of theearliest protostome and deuterostome metazoans and arean arena of active research. The Sellick Hill Formation ispart of the Normanville Group of the Stansbury Basin.This study considers the palaeontological, taphonomic,sedimentological and biostratigraphic implications of ahyolith rich, intraclast conglomerate horizon in the lowerSellick Hill Formation. Here, three new coelosclerito-phorans are described, as well as the validity of previouslydescribed specimens, since these fossils share no resem-blance with those previously described. The seeminglyunique nature of these fossils, although from one horizon,may have dire biostratigraphic implications and arediscussed in terms of their intra- and inter-basin andinternational correlations. The diagenetic and taphonomichistory is also discussed in an attempt to understand theenvironment of deposition and post depositional historyof this horizon. Here up to 14 diagenetic and taphonomicstages have been identified. The disarticulated and highlydamaged nature of recovered fossils, in addition to thesedimentology of this horizon implies a thanatocoenesicassemblage accumulated under periodic storm activity.The genesis of these seemingly unique fossils is, however,unknown. It is suggested that these sclerites were broughtinto the Normanville Basin from elsewhere under aperiod of storm activity. This study has also attempted toplace the biota and lower Sellick Hill Formation within thecontext of earliest Cambrian time. Based on the sum ofall data from the Sellick Hill Formation, in addition todescribed specimens, an Atdabanian, if not earlier, age issuggested for the Formation as opposed to the currentlyaccepted Botoman age.

Basin analysis, palaeontology and biostratigraphyof the Early to Middle Devonian Bokkeveld Group(Cape Supergroup), Western Cape(Poster)

Penn-Clarke, Cameron1; Rubidge, Bruce1; Jinnah, Zubair1;Almond, John2

1 Bernard Price Institute for Palaeontological Research, University of theWitwatersrand, Johannesburg, South [email protected]

2 Natura Viva CC, P.O. Box 12410 Mill Street, Cape Town, 8010 South Africa

Worldwide the Devonian Period is marked by theen masse colonization and diversification of terrestrialembryophytes, the so-called ‘Devonian Explosion’. Bodyfossils, as well as ichnofossils indicate that by the LateDevonian to Early Carboniferous, fully terrestrial-adapted arthropods (the arachnids, isopods, myriapodsand hexapods), as well as tetrapods, were adapted toliving on land. In the oceans, fish also underwent radia-tion with many ancient orders existing concomitantlywith more derived orders, in particular actinopterygiianand sarcopterygiian fish as well as true sharks (Neoselachii).Ammonoid cephalopods also made their first appearanceduring the Devonian Period. From palaeontologicalevidence, rocks of the South African Bokkeveld Group(Cape Supergroup) are considered to have been depos-ited between the Early to Middle Devonian. Consideringthe aerial extent of the Group, relatively little research hasbeen undertaken on the sedimentology, stratigraphy and

palaeontology of this very important aspect of the SouthAfrican fossil record. The rocks of the Bokkeveld Groupcontain a rich fossil fauna of Malvinokaffric Realm inver-tebrates, fish and plants. The Bokkeveld Group recordssedimentation as a series of five upward coarsening mega-cycles. Each megacycle is composed of a basal horizon-tally-bedded mudstone-siltstone. Lenticular and wavybedding is also present. These units grade upwards intoan overlying amalgamated sandstone body with a lateral(along strike) extent of >150 km. Initiation of each mega-cycle is related to a combination of reduction in sedimentsupply and basin deepening, most likely from extensionalsubsidence. Overall the Bokkeveld Group is a shallowingupward sequence with the overlying, conformable, andincreasingly terrestrial Witteberg Group. The pre-cursorsand events leading up to potential terrestrial environ-ments in the Witteberg Group, although hinted at by pre-vious work done on the Bokkeveld Group, have not yetbeen recognized and remain a field of potential study.This project aims to reassess the sedimentology of theBokkeveld Group, in addition to discussing lithologicalrelationships within a sequence stratigraphic frameworkin order to understand dynamics of the Cape Basin at timeof sedimentation at a section in the Cederberg Range,Western Cape. Fossil assemblages of the BokkeveldGroup too shall be addressed as well as any temporal orspatial controls over their occurrences/ranges in order todiscuss any environmental and/or ecologic controls pres-ent during this time frame.

A reconstruction of the sphenophyte Trizygiaspeciosa from the latest Permian WapadsbergPass locality in the Eastern Cape

Prevec, Rose1; Looy, Cindy2; Gastaldo, Robert3

1 Geology Department, Rhodes University, and Albany Museum, Grahamstown,South [email protected]

2 Department of Integrative Biology and Museum of Paleontology, University ofCalifornia, Berkeley, U.S.A.

3 Department of Geology, Colby College, Waterville, U.S.A.

The Wapadsberg Pass plant fossil locality, Eastern CapeProvince, has produced one of the youngest Glossopteris-dominated floras in the world, situated 70 m below what isgenerally considered to be the Permian/Triassic boundaryon the basis of regional vertebrate biostratigraphy.Although of unusually low diversity for a glossopteridflora, the quality of impression fossil preservation ishigh in this autochthonous deposit that is preserved in apossible reworked ash. This has resulted in the preserva-tion of very delicate structures including over 40 conesattributed here to the sphenopsid Trizygia speciosa.Although this sphenophyte is widespread across Gond-wana (it was first described from India by Royle in 1839),fertile organs have not been reported previously. Thecones are mostly dispersed, but several are attached tostems typical of Trizygia, that are gracile with expandednodes. This differs from the broader forms of Para-calamites that can be attributed to Phyllotheca australis, theonly other, fairly rare, sphenophyte found at the locality.Spores have been isolated from compression materialwithin cones. Although known from a number of Late

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Permian (Lopingian) localities in South Africa, well-preserved specimens of T. speciosa are rare, typically withonly one or a few consecutive leaf whorls preserved.However, dense mats of stems with multiple leaf whorlsare present at the Old Wapadsberg Pass. Examination ofthis unusual fossil flora has provided insights into themorphology of a little-understood sphenopsid, as well asproviding a glimpse of life prior to the greatest extinctionevent in Earth’s history.

Bone histology of the stegosaur, Kentrosaurusaethiopicus (Ornithischia: Thyreophora), from theUpper Jurassic of Tanzania

Redelstorff, Ragna1; Hübner, Tom2;Chinsamy-Turan, Anusuya1

1 Department of Zoology, University of Cape Town, Rondebosch, Cape [email protected]

2 Niedersächsisches Landesmuseum Hannover, Hannover, Germany

Using bone histology, a slow growth rate, atypical ofmost dinosaurs, has been interpreted for the highly derivedstegosaur, Stegosaurus (Ornithischia: Thyreophora), andthe basal thyreophoran Scutellosaurus (Redelstorff &Sander 2009; Padian et al. 2004). As the Thyreophora arebasal within the Dinosauria, this growth rate may reflect aplesiomorphic growth pattern within the clade and hasdirect implications for their biology. It is therefore impera-tive that further analyses of Thyreophoran bone histol-ogy are conducted. To this end, we analysed the bonehistology of the stegosaur Kentrosaurus from the Early Ju-rassic deposits of the Tendaguru beds of Tanzania.Kentrosaurus is both intermediate in size and phylogeneticposition between Scutellosaurus and Stegosaurus. We stud-ied the morphology and dimensions of all theKentrosaurus femora at the Berlin Museum of Natural his-tory, and we were permitted to sample five femora forhistological analyses. Our histological samples ofKentrosaurus reflect an ontogenetic series from subadult toadult specimens. We found that primary bone inKentrosaurus consists mainly of highly vascularizedfibro-lamellar bone with reticular vascular channels. Inaddition to LAGs and annuli, distinctive changes in the or-ganization of vascularization occur in the compacta,which seem to depict a regular cyclicity. This variation invascularity may reflect annual climatic fluctuations. Thebone deposition and growth rate in Kentrosaurus is signifi-cantly faster than that observed in Stegosaurus andScutellosaurus. Considering that Stegosaurus is thelarger-sized of the two stegosaurs, this contrasts with thedeductions made previously that small dinosaurs haveslower growth rates than large dinosaurs (Padian et al.2004). Our analysis of Kentrosaurus suggests that the slowgrowth rate is, not a phylogenetic characteristic of theThyreophora but may rather indicate a secondarily re-duced growth rate in the highly derived Stegosaurus.

First biarmoscuchian therapsid from thePristerognathus Assemblage Zone of theKaroo Basin, South Africa

Rubidge, Bruce; Day, Mike; Abdala, FernandoBPI Palaeontology, School for Geosciences, University of the Witwatersrand,

Johannesburg, South Africa

[email protected] / [email protected] /[email protected]

The Biarmosuchia, previously known from only a fewpoorly preserved specimens from South Africa and Russia(Sigogneau-Russell 1989), are considered to be the mostbasal therapsid clade and have recently been the subjectof renewed investigation with the description of severalnew genera (e.g. Rubidge & Kitching 2003; Rubidge et al.2006; Sidor & Rubidge 2006; Sidor et al. 2004; Sidor & Smith2007; Smith et al. 2006). Most of these new genera belong tothe Burnetiamorpha, a bizarre and apparently derivedclade of the biarmosuchia characterized by pachyostoticbosses on the skull roof. This group now comprises tengenera including Lemurosaurus, Lobalopex, Lophorhinus,Paraburnetia , Burnetia , Niuksenitia , Bullacepahlus ,Pachydectes and a new undescribed form from Malawi.Apart from Lemurosaurus, which comprises two speci-mens, the rest of the genera are known from only a singleholotype specimen. Two of these genera are from Russia,one from Malawi, and eight are from South Africa.

Burnetiamorphs have a Mid-Late Permian stratigraphicrange and in South Africa are known from all the Permianbiozones of the Beaufort Group, save the Eodicynodonand Pristerognathus Assemblage Zones (Sidor & Smith2007), with some of these biozones having more than onegenus. Previous phylogenetic analyses have linkedPachydectes and Bullacephalus, the stratigraphically lowestoccurring genera, with Burnetia, the burnetiamorph fromthe latest Permian of South Africa; such a result suggeststhat most of the biarmosuchian lineage is unrecorded.This paper describes a new specimen, the first from thePristerognathus Assemblage Zone, which adds to ourunderstanding of the taxonomic diversity and biostrati-graphic distribution of the taxon.

Rubidge, B.S. & Kitching, J.W. 2003. A new burnetiamorph (Therap-sida: Biarmosuchia) from the lower Beaufort Group of South Africa.Palaeontology 46, 199–210.

Sidor, C.A., Hopson J.A. & Keyser, A. 2004. A new burnetiamorph(Therapsida: Biarmosuchia) from the Teekloof Formation, Permian ofSouth Africa. Journal of Vertebrate Paleontology 24, 938–950.

Sidor, C.A. & Rubidge, B.S. 2006. A new biarmosuchian (Therapsida:Biarmosuchia) from the Beaufort Group of South Africa pp. 76–113. In:Carrano, M.T., Gaudin, T., Blob, R. & Wible, J. (eds), Amniote Paleo-biology: Perspectives on the Evolution of Mammals, Birds, and Reptiles.Chicago, University of Chicago Press.

Sidor, C.A. & Smith, R.M.H. 2007. A second burnetiamorph therapsidfrom the Permian Teekloof Formation of South Africa and itsassociated fauna. Journal of Vertebrate Paleontology 27, 420–430.

Sidor, C.A. & Welman, J. 2003. A second specimen of Lemurosaurus pricei(Therapsida: Burnetiamorpha). Journal of Vertebrate Paleontology 23,631–642.

Sigogneau-Russell, D. 1989. Theriodontia I. In: Wellnhofer, P. (ed.), Ency-clopedia of Paleoherpetology, Part 17B, 1–127. Stuttgart, Gustav Fischer.

Smith, R.M.H., Rubidge, B.S. & Sidor, C.A. 2006. A new burnetiid(Therapsida: Biarmosuchia) from the Upper Permian of South Africaand its biogeographic implications. Journal of Vertebrate Paleontology 26,331–343.

The gigantism in fossil amphibians

Sanchez, SophieUppsala University, Sweden

[email protected]

The appearance of large, even giant, animals in theevolutionary history of vertebrates has long raised theattention of biologists and paleontologists, especially

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when regarding fishes, marine reptiles, dinosaurs andmammals. Gigantism is defined as an excessive growthof the stature compared to the mean size of the adultindividuals of the same clade which is equal to or morethan two standard deviations. During their evolution,amphibians (as non-amniotic tetrapods) also exhibitedgiant body lengths, attaining up to six or seven metresduring the Triassic period. Understanding how such giantsizes could have been reached in fossil amphibians is ofmajor interest, especially when regarding the size distri-bution of the lissamphibians (1.8 m for the largerlissamphibian: Andrias davidianus, the so-called giantChinese salamander). In order to find some explanationsfor this gigantism in fossil amphibians, I have performed apreliminary histological study on the long-bone mid-shafts of fourteen species among the various clades of theTemnospondyli, Seymouriamorpha and Embolomeri. Itseems that gigantism in fossil amphibians may have beena hyperpituitary gigantism observable at the histologicallevel, i.e. a combination of 1) an increase of growth rates(which can be 50–80 times higher in Wetlugasaurusangustifrons than in Apateon pedestris, for example), assuggested in this study by skeletochronology; and 2) ablocking of the development suggested by the retentionof juvenile histological features in adults (=paedomor-phosis). These histological phenomena could certainly belinked to environmental conditions; increases in size andhypothyroid activity have already been observed in tetra-pods, which have secondarily returned to the aquaticenvironment. It is now necessary to focus on the ecologi-cal changes, which occurred during the Permian-Triassiccrisis, in order to better understand the physiological reac-tions of these giant fossil amphibians, especially amongtemnospondyls, which lived up to the Early Cretaceous.

The global biogeography, taxonomyand chronological distribution ofinsular proboscideans(Poster)

Scarborough, Matthew E.Department of Zoology, University of Cape Town, Rondebosch, South Africa

[email protected]

Proboscidean fossils are known from many islandsaround the world, including many of the Mediterraneanand southeast Asian islands, dating from the Miocene toMid-Holocene. A checklist of the global occurrence ofproboscideans on islands is presented, and the chronology,taxonomy and presence of size-reduction in insularendemics is reviewed. The greatest diversity in insularproboscideans occurs in the genera Palaeoloxon, Stegodonand Mammuthus during the Pleistocene, all of whichrepeatedly evolved endemic species on islands. Homo-plastic traits in endemic proboscideans and their possibleheterochronic evolutionary mechanisms are discussed.

Fluctuating Miocene climates and ecosystems atthe southern tip of Africa: a multiproxy approach

Sciscio, Lara1; Neumann, Frank H.2; Roberts, David3;Tsikos, Hari1; Scott, Louis4; Bamford, Marion5

1 Department of Geology, Rhodes University, Grahamstown, South [email protected]

2 Forschungsstelle Palaeobotanik, University of Münster, Germany

3 Council for Geoscience, Bellville, South Africa4 Department of Plant Sciences, University of the Free State, Bloemfontein,

South Africa5 BPI Palaeontology, School of Geosciences, University of the Witwatersrand,

Johannesburg, South Africa

The largest Lagerstätte of excellently preserved Mio-Pliocene terrestrial fauna in South Africa is located atLangebaanweg (LBW), now the West Coast Fossil Parksituated ~120 km north of Cape Town. This research dealswith a core drilled at Langebaanweg ‘E’ Quarry, whichcaptures Miocene fluvial deposits of the ElandsfontynFormation (Sandveld Group), unconformably underlyingthe Mio-Pliocene faunal deposits. The work aims to con-strain fluctuations in climate and ecosystems in the regionduring the Miocene, using palynological, biogeochemical,stable isotope (C, N) and sedimentological approaches. Anovel proxy for the reconstruction of terrestrial palaeo-climates, which examines branched glycerol dialkyl glyc-erol tetraether (GDGTs) membrane lipids produced bysoil bacteria, has been used. We have applied thesebiomarkers (or molecular ‘fossils’) in conjunction withpalynology to the organic sediments of the poorly studiedElandsfontyn Formation.

The mean annual temperature (MAT) and ambient pHvalues of the organic-rich horizons, at time of deposition,were calculated using the Methylation index of branchedtetraethers (MBT) and cyclization ratio of branchedtetraethers (CBT) proxies. LBW samples, which span17–33 m below sea level, had MATs ranging between12.4°C to 26.6°C and pH range from 4.4 to 6.4 over the 16 mof core studied. Furthermore, samples showed a narrow,light (from –25.52‰ to –24.27‰) Ç

13C value distributionand overall low C/N ratios. Palynological investigationsupplemented earlier studies, reaffirming alternatingsequences of tropical and subtropical elements andcomplementing calculated MAT results. Pollen resultsfrom the subsection 1 (30.89–33 m) provide evidence of aspecies rich Podocarpus-dominated forest with MATs at16.2°C. The pollen-bearing section between 16.45 m and20.35 m depth (subsection 2) shows initially less humidconditions, with MATs between 15.4–26.6°C, similar to thebottom of the section with high podocarp percentages,low Restionaceae and aquatics. The results suggest thatthe regional Miocene climate showed high amplitudefluctuations (possibly driven by orbital forcing as seen inmarine cores), underscoring the potential of biogeo-chemistry for unravelling past climates and ecosystems.

Early Jurassic trace fossil and conchostracan-bearing lacustrine deposits: refining the ancientdryland ecosystem of the Upper Elliot Formation(Karoo Supergroup) in Lesotho(Poster)

Sciscio, Lara1; Bordy, Emese M.1; Knoll, Fabien2;de Cock, Michiel3

1 Department of Geological Sciences, University of Cape Town, Rondebosch,South [email protected] / [email protected] /

2Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, Madrid,[email protected]

3 Department of Geology, University of Johannesburg, South [email protected]

The Elliot Formation (Late Triassic – Early Jurassic) of the

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Karoo Supergroup is a fossiliferous, fluvio-lacustrine redbed succession that is believed to capture the end-Triassicmass extinction event and subsequent biotic recovery inthe Early Jurassic of southern Africa. A new study site insouth-western part of Lesotho (Ha Seeiso, near Matelile,Mafeteng District) reveals a previously unreportedlacustrine deposit within the upper Elliot Formation.This interbedded succession of fine-grained sandstonesand mudstones is ~17 m thick, and is characterized bylaterally persistent, tabular beds that show no evidencefor channelling. The grain sizes, sedimentary structures,facies architecture and invertebrate and vertebrateichnofossil assemblage suggest energy level fluctuationsin an otherwise low energy depositional setting. Further-more blue-black, finely laminated organic-rich mudstoneswhich lack any pedogenic alteration features imply a lakesystem with multiple anoxic periods. These unique organicunits also contain conchostracan (Cyzicus) and ostracodvalve impressions which suggest seasonal evaporationevents. Ostracod-bearing beds have only previously beenmentioned in the Clarens Formation. This new fossillocality encapsulates the life and events of an ephemerallake in the arid depositional environment of the upper-most Elliot Formation. Further geochemical and palyno-logical examinations of the organic units are being under-taken in order to improve our understating of the dynam-ics of this Early Jurassic lacustrine ecosystem.

A new method to determine volume of spiralbromalites(Poster)

Shelton, Christen D.Steinmann Institut of Geology, Mineralogy and Palaeontology,

University of Bonn, [email protected]

Lower Permian vertebrates from both terrestrial andaquatic organisms have been collected from ArcherCounty, Texas, U.S.A. for over a century. This includespreserved shark cartilage and spiral bromalites presumedto have been produced by freshwater sharks of the genusOrthacanthus. Specimens were collected in the newlynamed Archer City Bone Bed V (Archer City Formation).Physical characteristics (length, width, height, mass,eccentricity, volume, and density) were measured andrecorded for 300 spiral heteropolar bromalites and com-pared by linear regression analysis. The spiral bromaliteshad an average length of 31.74 mm; width 14.43 mm;height 10.51 mm, and mass 7.971 g. Eccentricity aroundthe longest axis was between 0.9399 and 0.0602. In addi-tion to the statistical collection, a new formula is proposedto help determine the volume of the elliptical fossils whenvolume determination by water displacement is not anoption. The regression analysis between the observedvolumes obtained by water displacement and thecalculated volumes showed R2 = 0.9836. The percentagedifference between the two measurements ranged from0.04% to 56% (95% of the statistical sample had a percent-age difference of <30). Data from this study and theformula developed here will help to understand not onlythe spiral bromalites from this area, but also bromalitesfrom different localities, geological ages, and different

origins. Problems with this formula include accuracy ofthe physical measurements, which would be skeweddue to any extra matrix on the specimen. Also, the formulaassumes that all specimens terminate in a point, but manybromalites are incomplete or have blunt tips, due to de-structive forces of nature, erosion, or lack of preservation.This would allow some room for error and perhapsexplain the extreme percentage difference in some speci-mens. These data, if compiled with data from the analy-sis of specimens from other localities, may ultimatelyreveal size groupings that could reflect the presence ofpreviously unknown taxa that possessed a spiral valve andcoexisted with the Orhtacanthus sharks. From the data wecan only conclude that a single species produced theheteropolar bromalites at this bone bed. A standard prece-dent for classifying these fossils based on external as wellas internal morphometric data is crucial to understandingtheir origin and producers.

Ophiacodon long bone histology – testing theBrinkman hypothesis

Shelton, Christen D.; Sander, P. MartinSteinmann Institute of Geology, Mineralogy and Palaeontology,

University of Bonn, [email protected]

The validity of species in a genus that only differ in sizehas always come into question. Histological analysis is oneof the most definitive methods of species determinationbased on ontogenetic stages of the diaphyseal mid-shaftof long bones, usually femora or humeri, which areknown to contain the best record of growth. Innon-therapsid synapsids (pelycosaurs), some specieshave been identified solely on the basis of size regardlessof the degree of ossification observed in the epiphysis.Brinkman (1988) attempted to overcome this problem byestablishing size-independent morphogical criteria onputative growth series of pelycosaurs including Ophiacodon.Based on degree of ossification of the epiphyses and thedevelopment of specific anatomical structures, he subdi-vided the putative growth series into five stages. Here wetest his hypothetical growth series using histologicalindicators. The growth series comes from the RattlesnakeCanyon locality (Nocona Formation, Lower Permian) inArcher County, Texas, U.S.A.. We conclude that the histol-ogy of the Ophiacodon humerus growth series of Brinkmandoes support the morphological ontogenetic stages heassigned to those specimens. The general histology ofthe humeri consist of a cortical bone matrix made up ofparallel-fibred and woven bone and is highly vascularizedby radial canals with varying degrees of lamellar boneinfilling. This gives the cortex a ’spoked bicycle wheel’ ap-pearance. The medullary cavity is filled with a network oftrabecular bone. The outer most cortex of the largesthumerus, assigned to stage five by Brinkman, contains anexternal fundamental system (EFS) and a sharp decreasein vascularization. This unequivocally indicates that theanimal had passed sexual maturity. However, the timerepresented by the EFS is unknown. The smallest humerusexamined was determined by Brinkman to be a stage twoand its histology preserves the neonatal line in the deepcortex. This is an indicator of the time when the animal

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presumably hatched, and is the boundary between origi-nal embryonic bone and tissue deposited postnatally. Thehistology and vascularization of the cortex is very similarto that seen in Dimetrodon natalis and, surprisingly, basalsauropterygians. This tissue has been named incipientfibro-lamellar bone. It is the precursor to the true fibro-lamellar bone seen later in therapsids and mammals.

Field evidence for stepped mass extinction in theSouth African Karoo Basin at the end-Permian

Smith, Roger1; Botha-Brink, Jennifer2

1 Iziko South African Museum, Cape Town, South [email protected]

2 Karoo Palaeontology, National Museum, Bloemfontein, South Africa

The southern Karoo Basin of South Africa contains anuninterrupted terrestrial record of the Permo-Triassicboundary (PTB). Isotope- and magneto-stratigraphyconfirm that these fluvial strata are approximately thesame age as zircon-dated marine PTB sections (252 Ma).To date, our team has found 580 identifiable, in situ verte-brate fossils, mostly therapsids, in PTB exposures at fourseparate locations. Biostratigraphic range plots reveal apronounced extinction event within the same strati-graphic interval in each of the PTB sections as well as aregular sequence of taxon disappearances and tapho-nomic signatures that are interpreted as reflecting realchanges in the original populations due to rapid climaticdrying and drought. Within the uppermost 45 metres ofthe Balfour Formation, the sequence of sedimentologicalfacies reflects progressive straightening of the channelsand floodplain rubification. Although thicknesses vary,this sequence remains consistent across all sections and isinterpreted as indicative of a rapid climatic warming anddrying with the onset of a monsoon-type rainfall regime.In the lowlands of the Karoo basin ground-level ferns,clubmosses and liverworts were the most susceptible tothe lowering of groundwater. Consequently the formerlyflourishing populations of small gregarious herbivorousdicynodonts such as Diictodon and Pristerodon decreasedand eventually disappeared along with their medium-sizedgorgonopsian predators (Aelurognathus, Cyonosaurus). Asdroughts became more frequent the Glossopteris riparianwoodlands thinned out causing large-bodied dicyno-donts (Dicynodon, Aulacephalodon, Dinanomodon) and theirattendant carnivores to die off. A terminal drought phaseis recognizable in all the sections as a roughly 5 metre-thick interval containing 1–3 beds of generally rubifiedfinely laminated mudrocks with very little pedogenicmodification. Rare skulls and fragmental caniniform pro-cesses of the larger Permian taxa L. maccaigi, Moschorhinusand a very rare skull of Dicynodontoides occur in this inter-val, the top of which is regarded as a reliable litho-stratigraphic marker of end of the End-Permian massextinction in the Karoo Basin.

The dietary behaviour of mammals fromCooper’s Cave

Steininger, ChristineInstitute of Human Evolution, School of Geosciences, University of the

Witwatersrand, Johannesburg, South [email protected]

Preliminary ecological reconstructions based on com-

parisons of fossil and modern mammals suggests this siteis a predominately a grassland environment (deRuiteret al. 2009). This argument is flawed, as the behaviour of amammal in the past may not be the same as the present.For example, the modern impala is considered a savannaantelope that consumes a mixture of grass and leaves;however, using a direct method, stable carbon isotopeanalysis, this same species fed primarily on tree leaves inthe past. In addition, based on its dietary preference wecan infer that this species may have favoured a woodlandenvironment. Research conducted for my doctoral disser-tation based on the dietary behaviour of several bovidspecies from Cooper’s Cave (locality D) and Swartkrans(Members 1–3) suggests that, while grass was a compo-nent of the landscape, woody vegetation predominated,at least in the immediate vicinity of these caves. Researchis under way to sample the dietary behaviour of a largercomponent of the mammal community from Cooper’sCave. This site has a diversity of mammals, includinghominins and early stone tools with uranium series datesof 1.5–1.4 Ma. Two questions are addressed: (1) will arepresentative sample of diverse mammals from Cooper’sCave show a predominately-woody vegetation structureand (2) How are mammals including hominins parti-tioned within this environment based on their dietarybehaviour?

Tooth root morphology in the giant African bearAgriotherium africanum (Mammalia, Carnivora,Ursidae) and its implications for feeding ecology

Stynder, Deano1; Kupczik, Kornelius2

1 Department of Archaeology, Faculty of Science, University of Cape Town,Rondebosch, South [email protected]

2 Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum,Universität Jena, Jena, Germany

Tooth root surface areas serve as a proxy for bite forcepotentials, and by extension, dietary specialization inextant carnivorans. In this study, we investigate the feed-ing ecology of the extinct ursid Agriotherium africanum bycomparing its root surface areas (reconstructed with theaid of computed tomography and three-dimensionalimage processing) and bite force estimates, with those ofextant carnivorans. Our results show that in absoluteterms, A. africanum exhibited higher canine and carnassialbite forces and root surface areas than the extantcarnivorans used in this study. However, when adjustedfor skull size, A. africanum’s canine roots were smaller thanthose of extant large predators. As teeth are the limitingfactor in the masticatory system, low canine root surfaceareas suggest that A. africanum struggled to bring downlarge terrestrial prey. In addition, its adjusted carnassialroot sizes were found to be smaller than those of extanthard and tough object feeders, but larger than those ofextant mesocarnivorous ursids and Ursus maritimus. Thefact that it also possessed carnassials with hyaena-like(pointed) cusps and exhibited its highest post-canine rootsurface areas in the carnassial region, suggest thatA. africanum was more competent at scavenging thanextant ursids. As the largest carnivore on the early Plio-cene African landscape there was an ecological opportu-

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nity for it to develop into a specialist scavenger of largeterrestrial vertebrates, a niche that it likely occupied givenits dental root morphology. Be this as it may, the presenceof premasseteric fossae on its mandible suggests that itwould have consumed plant material on occasions.

Piltdown Man and Teilhard de Chardin

Thackeray, FrancisInstitute of Human Evolution, School of Geosciences, University of the

Witwatersrand, Johannesburg, South [email protected]

On January 5, 1913, the French palaeontologist, philoso-pher and priest, Pierre Teilhard de Chardin, wrote anessay in a French Jesuit journal called Etudes. The essaywas written within a few weeks of the official announce-ment in London of the discovery of ’Piltdown Man’,assigned to Eoanthropus, which included a human craniumand a carefully modified mandible and isolated canine ofan orangutan. The opening sentence of Teilhard’s essay isas follows: ’Il fut un temps où la préhistoire méritait d’êtresuspectée ou plaisantée’. This can be translated to mean’There was a time when the study of prehistory deservedto be suspected, or deserved to be the subject of jokes’.Attention is drawn to the mere possibility that almostimmediately after the official announcement of ’PiltdownMan’ in Burlington House in London (December 18,1912), Teilhard wrote this sentence in Paris when he mayhave been disturbed by the fact that English palae-ontologists, including Smith Woodward of the British Mu-seum (Natural History), may have been ’taken in’ by ajoke. If Teilhard had known even before January 1913 thatPiltdown was intended as a joke, he could have beendeeply concerned by the potential adverse impact if the’Piltdown joke’ was taken seriously. It is suggested that hewas writing with Piltdown in mind when he wrote theopening sentence of his essay in January 1913, referring tothe study of prehistory, including palaeo-anthropology,as a field which deserved to be ’suspected’ and whichdeserved to be a ’subject of jokes’. Within the Jesuit order,it was considered acceptable to lie provided it was a joke(Thackeray 2012). ’Piltdown Man’, announced 100 yearsago, is likely to have been a joke associated not only withTeilhard de Chardin but also Martin Hinton and CharlesDawson. The latter was known to have been a perpetratorof forgeries, and is also likely to have been involved in Pilt-down (perhaps in the case of ’Piltdown II’).

New data on the cranial anatomy,phylogenetic position and biogeographyof Bunostegos akokanensis

Tsuji, Linda1; Sidor, Christian1; Smith, Roger2;Steyer, Jean-Sébastien3

1 Burke Museum and Department of Biology, University of Washington,Seattle, [email protected]

2 Iziko South African Museum, Cape Town, South Africa3 Department Earth History, CNRS-Museum National d’Histoire Naturelle,

Paris, France

Newly recovered cranial material of Bunostegos akokanensis,a pareiasaurian reptile known only from the Upper Perm-ian Moradi Formation of northern Niger, permits aredescription of the taxon in addition to inclusion of new

information in a phylogenetic analysis of pareiasauro-morphs. Bunostegos is highly autapomorphic, with diag-nostic cranial features including: two or three hemisphericalbosses located at the anterior end of the snout; elongate,laterally projecting supraorbital ‘horn’ formed by anenlarged postfrontal; large foramen on ventral surface ofpostfrontal; hemispherical supratemporal boss atposterolateral corner of skull roof. We addressed thephylogenetic position of Bunostegos by incorporating itinto a cladistic analysis of 29 parareptilian taxa (includingall 21 currently valid named pareiasaurs) and 127 cranialand postcranial characters. The results place Bunostegos asmore derived than the South African Middle Permianforms such as Bradysaurus and Embrithosaurus, and as thesister taxon to the Upper Permian taxa including theRussian Deltavjatia plus Velosauria. Characters related tothe cranial sculpture and to the size and placement of thetabulars appear to be similar to more derived pareiasaurssuch as Elginia from Scotland and Arganaceras fromMorocco, but parsimony indicates that these featuresappeared independently in Bunostegos. The relationshipsof velosaurian pareiasaurs, including Anthodon, Pumilio-pareia, and Scutosaurus, were consistent with those ofprevious analyses.

Pareiasaurs are important biostratigraphic markerswithin the five Permian assemblage zones established forSouth Africa’s Beaufort Group. In addition, the occur-rence of Pareiasuchus and Anthodon in the Ruhuhu Basin ofTanzania and Pareiasuchus in the Luangwa Basin ofZambia has contributed to the regional correlation ofthese strata. The tetrapod fauna of the Moradi Formation,however, has proven difficult to correlate to other Africanassemblages because it contains endemic genera (viz.Bunostegos, Moradisaurus, Nigerpeton, and Saharastega).The single therapsid known from Niger, an indeterminategorgonopsid, is suggestive of an Upper Permian assign-ment, but is hardly definitive. Moreover, the lack of bothdicynodont herbivores and Glossopteris in the Permian ofNiger indicates a markedly different community structureand supports the theory that central Pangaea wasbiogeographically isolated from the rest of the super-continent by desert-like conditions. Our improved under-standing of the morphology and phylogenetic positionof Bunostegos helps to piece together a more completepicture of Permian tetrapod evolution.

Post-mortem in 3D: new methods to assess thespatial taphonomy of the early hominins from theMalapa site

Val, Aurore1,2; Backwell, Lucinda R.1; Berger, Lee3;d’Errico, Francesco2,4

1 Bernard Price Institute for Palaeontological Research, School of Geosciences,University of the Witwatersrand, Johannesburg, South [email protected] / [email protected]

2 PACEA UMR 5199-CNRS, Université Bordeaux 1, France3 Institute for Human Evolution, University of the Witwatersrand, Johannesburg,

South [email protected]

4 Department of Archaeology, History, Cultural Studies and Religion, University ofBergen, Postboks 7805, NO-5020 Bergen, [email protected]

Two nearly complete skeletons of early hominins

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(Australopithecus sediba; Berger et al. 2010) were found atthe Malapa cave site (Bloubank Valley, Gauteng Province,South Africa), in a deposit dated at 1.977 Ma (Dirks et al.2010; Pickering et al. 2011). Some anatomical elementswere still in articulation and their bone surface shows agood state of preservation. This case scenario (highnumber of remains per individual, preservation of somearticulations, good preservation of the outer bone mor-phology, high percentage of complete bones) is extremelyrare in the context of early hominin discoveries. In orderto understand the fossilization processes that have led tosuch an unusual level of preservation, a comprehensivetaphonomical analysis of the hominins and associatedfaunal remains is currently being conducted. We summa-rize here the techniques and methods used to investigatethe taphonomic implications of the spatial arrangementand orientation of the hominin remains.

The remains of two near-complete individuals (MalapaHominid 1, a juvenile male and Malapa Hominid 2, anadult female) include 183 specimens, 46 of which wererecovered in situ and 137 in various ex situ blocks of clasticcalcified sediment. Each phase of the excavation andpreparation processes, including the spatial coordinatesof the in situ remains, was recorded. By combining thisinformation and the 3D virtual renderings of each speci-men to geological and sedimentary data, we are able topropose a best-fit 3D model of the original position of thetwo A. sediba individuals within the deposit, prior tomining damage. This model has been created by applyingCT and micro-CT scanning techniques and 3D renderingsoftware (Avizo 6.3).

Berger, L.R., de Ruiter, D.J., Churchill, S.E., Schmid, P., Carlson, K.J.,Dirks, P.H.G.M. & Kibii, J.M. 2010. Australopithecus sediba: a new speciesof Homo-like australopith from South Africa. Science 328, 195–204.

Dirks, P.H.G.M., Kibii, J.M., Kuhn, B.F., Steininger, C. Churchill, S.E.,Kramers, J.D., Pickering, R., Farber, D.L., Meriaux, A-S., Herries, A.I.R.,King, G.C.P. & Berger, L.R. 2010. Geological setting and age ofAustralopithecus sediba from southern Africa. Science 328, 205–208.

Pickering, R., Dirks, P.H.G.M., Jinnah, Z., de Ruiter, D.J., Churchill, S.E.,Herries, A.I.R., Woodhead, J.D., Hellstrom, J.C. & Berger, L.R. 2011.Australopithecus sediba at 1.977 Ma and implications for the origins ofthe genus Homo. Science 333, 1421–1423.

Trackways of bipedal reptiles from Giant’s Castle,KwaZulu-Natal(Poster)

Van Dijk, D. EduardDepartment of Plant and Animal Biology, University of Stellenbosch, Stellenbosch

[email protected]

Among trackways in Lesotho described by Paul Ellen-berger in 1970 was a genus of reptiles, Molapopento-podiscus, with type species M. pilosus , which was consid-ered to be bipedal. A second species, M. supersaltator, fromGiant’s Castle, KwaZulu-Natal, was described by Ellen-berger from photographs sent to him. Each species wasillustrated by drawings of a pair of prints and indicationsof the length of a leap (hop). The upper and lower surfacesof a slab with M. supersaltator tracks was subsequentlyillustrated (Van Dijk 1978), making it quite clear that themode of locomotion had been correctly interpreted. In thesame publication was an illustration of the upper surfaceof a slab which included what were considered to be

prints of a biped with distinctly curved toes. In the ab-sence of a slab with a sequence of underprints whichcould be related to upper prints with curled toes, descrip-tion of a distinct taxon was deferred. A slab assembledfrom scree fragments that has a single pair of upper andlower prints, and another slab with both upper and lowerprints of one side of a short sequence has made it possibleto describe a second taxon

The bipedal prints have variously been attributed tomammals and pterosaurs. There is evidence to supportthe latter view.

Calcified wood from Mkuze, KwaZulu-Natal(Poster)

Van Dijk, D. Eduard1; Bamford, Marion2

1 Department of Plant and Animal Biology, University of Stellenbosch, [email protected]

2 BPI Palaeontology, School of Geosciences, University of the Witwatersrand,[email protected]

In the late 1950s staff at the entrance of Mkuze Reserve,KwaZulu-Natal, were aware of fossil tree trunks not farfrom the offices. One such trunk was photographed at thetime and loose pieces collected. The site has been identi-fied as Nhlohlela Hill and Swamp, now some distancefrom the present entrance to the reserve.

Large calcite crystals suggested that histological detailwas not retained. However, thin chips showed cellulardetail. When pieces were sectioned and section surfaceswere polished, and lightly etched with acid, it was foundpossible to detect detail in cellulose acetate peels. Groundsections were also made. Transverse sections showedsome variation in cell size and cell wall thickness, but treerings were not obvious. Tangential sections showednarrow groups of radial sections, mostly only one cellwide and about ten cells high. Repeated attempts atproducing radial sections as peels or sections have yieldedno useful ones.

In 2011 narrow regions that appeared to be iron oxidewere observed and sections along them were attempted.Bordered pits were observed as well as other microscopicdetails.

Origin of Early Triassic Lystrosaurus bonebedsfrom the Karoo Basin, South Africa: theirsignificance for faunal recovery in a post-extinction world

Viglietti, Pia A.1; Smith, Roger, M.H.2; Compton, John S.1

1 Department of Geological Sciences, University of Cape Town, Rondebosch,South [email protected] / [email protected]

2 Iziko South African Museum, Cape Town, South [email protected]

The most devastating mass-extinction event Earth hasexperienced occurred at the end of the Permian periodapproximately 252 Ma ago. The southern Karoo Basin isone of the few places in the world that preserves an almostcontinuous stratigraphic record of terrestrial sedimenta-tion through the Permian-Triassic Boundary (PTB).Emphasis on the rapid annihilation of marine andtetrapod faunas during the PTB extinction event hasgenerally overshadowed the nearly as rapid radiation and

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regeneration of fauna in earliest Triassic Karoo environ-ments. The expansion of mammal-like reptiles (therapsids)of the Lystrosaurus Assemblage Zone fauna into anincreasingly arid Karoo Basin is associated with a changein fossilization mode from isolated skulls in the latestPermian and bone material to articulated ’curled up’skeletons and multi-individual accumulations (bonebeds)in the early Triassic. Lack of epiphyses and skull sizessuggest only subadult Lystrosaurus are present inbonebeds. Clustered articulated ribs among disarticulatedmaterial and minor weathering of bones are evidence of aperiod of desiccation where bones lay on the surface forup to five years before burial (Botha & Smith 2005). Thelithology surrounding the bonebeds consists of floodplainmudrocks, carbonate nodules, and sand-filled mudcrackscapped by coarse sediments indicative of rapid depositionfollowing floods. Carbonate nodules were sampled andshowed Ç

13C values ranging from –8.5 to –5‰ and Ç18O

values of 13.5 to 16 . These values support the formation ofthe carbonate nodules under a cool semi-arid climate athigher latitude (~50°S) than the current Karoo Basin. Thisstudy agrees with previous studies arguing for the forma-tion of bonebeds during extended episodic periods ofdrought in the earliest Triassic Karoo Basin. The presenceof ichnofossils such as vertebrate burrow-fills suggeststhe use of burrows possibly by Lystrosaurus and otherfauna (such as cynodonts) to escape extreme environmen-tal conditions. Being able to adapt to these extreme condi-tions was key to the survival of certain species into theEarliest Triassic, but some species appeared to have beenbetter adapted than others. This is shown by thedisappearance of Lystrosaurus by the Middle Triassic andthe survival of the cynodonts, the ancestors of the mammallineage.

Anatomical fine structure of brain cases:application in international biostratigraphy

Welman, JohannUniversity of Limpopo, South Africa

[email protected]

In classical South African biostratigraphical studies,strata were characterized by the presence of index fossils.In more recent times, biostratigraphic characterization ofstrata has been done using the fossil assemblages and theranges of fossils that indicate boundaries of biostrati-graphic divisions.

In the international context, however, a species some-times occurs in different-aged faunal assemblages and inanomalous stratigraphic positions (i.e. in more than oneteilchron). These anomalous occurrences may be a functionof the time and space aspects of evolution, the longevity ofspecies and the time it took for a species to disperse geo-graphically over Pangaea. This creates questions aboutthe relative age of species in the different localities, sincethe mere presence or absence of the species apparentlyhas limited significance for refined terrestrial biostrati-graphy.

The introduction of international phylozones basedupon informed anatomical studies presents a way out ofthis dilemma. If the anatomy of closely related forms in

evolutionary lineages can be studied, trends in theirevolution can be determined and landmark develop-ments identified so that this succession of irreversibleanatomical evolutionary events can be used in chrono-stratigraphy. In this way refined functional-anatomicalstudies of the diversification of reptiles can pave the wayto a better determination of the biochronozones of terres-trial strata.

In the early diapsid reptile lineages leading up to birdsand dinosaurs, the fine structure of the brain cases of aseries of reptiles has been studied and the anatomicalchanges identified that gradually and irreversibly devel-oped between 290 and 150 Ma. Asuming the origin ofbirds from early archosauromorph reptiles, the origin offlight in an arboreal setting caused changes in the anatomyof the brain and ear that happened in a logical func-tional-morphological progression. The increase in thebrain size during the origin and early evolution of birds ischaracterized by the gradual co-evolution of otic andbasicranial structures. The presence or absence ofapomorphic structures that resulted from these evolu-tionary changes serve as precise indicators of relative agewithin the lineage composed of related species from local-ities internationally that could consequently serve as indi-cators of the relative age of the terrestrial sedimentaryrocks in which different taxa occur.

In the diverse dinosaur lineages and their Pangaeanarchosauromorph ancestors, the gradual evolution of thecomplex median Eustachian system driven either bythermoregulation or pressure equilization as possiblefunctional impetus, gave rise to well-defined clades withspecific landmark anatomical apomorphies. These devel-opments open up the possibility of high resolution strati-graphic correlation of the Lower Triassic to Cretaceousaquatic and terrestrial sediments internationally.

The Olduvai Hominid 8 (OH 8) foot and the statusof Homo habilis: Where do we stand?

Zipfel, Bernhard1; DeSilva, Jeremy2; Van Arsdale, Adam3;Tocheri, Matthew4; Weiss, Elizabeth5

1 Institute of Human Evolution, School of Geosciences, University of theWitwatersrand, Private Bag 3, Johannesburg, 2050 South [email protected]

2 Anthropology Department, Boston University, 232 Bay State Road,Boston, MA 02215, U.S.A.

3 Department of Anthropology, Wellesley College, Wellesley, MA, U.S.A.4 Human Origins Program, Department of Anthropology, National Museum of

Natural History, Smithsonian Institution, Washington D.C. 20013-7012, U.S.A.5 Department of Anthropology, San Jose State University, One Washington

Square, San Jose, CA 95192-0113, U.S.A.

The OH 8 fossil foot assemblage has been important forinterpreting foot evolution and the evolution of homininlocomotion. It is generally accepted that OH 8 belongs to asingle bipedal individual but debates, both past and pres-ent, are centred on maturity and taxonomic attribution. Ithas been suggested by some, that the presence of what areperceived as unfused metatarsal epiphyses, that OH 8 wasa juvenile, associated with the Homo habilis type OH 7. Ithas also been associated with the OH 35 tibia and fibula,but this has been refuted. Accurately assessing the age ofthe OH 8 individual at death is important for interpreting

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the functional morphology of these fossils and the loco-motion practiced by early Pleistocene hominins. Here, wereview the debate around the juvenile and adult status ofthe OH 8 foot and present evidence that strongly suggeststhat the OH 8 assemblage is that of an adult. A study of thepatterns of epiphyseal fusion of the original OH 8 fossils,subadult feet from modern humans, chimpanzees, andgorillas suggest that the original interpretation of the OH8 foot as an adult individual may be the correct one. Basedon the timing of epiphyseal fusion in the ape and humanfoot, if the OH 8 foot is from a juvenile with unfused meta-tarsal heads, the epiphysis of the first metatarsal baseshould also be unfused. However, the epiphysis of thefirst metatarsal of OH 8 is fully obliterated and none of thesample of feet studied shared this pattern of metatarsal

fusion. A radiographically visible epiphyseal scar on theOH 8 first metatarsal has been used as evidence for recentfusion and thus subadult status. To this end, a radio-graphic study of first metatarsals of humans from ages 17to 88 years was carried out. The results show that the basalepiphyseal line is visible past the age of epiphyseal fusionin 29% of the radiographed first metatarsals. Statistically,there was no relationship between the loss of the epi-physeal scar and age. The presence of the epiphyseal scartherefore does not necessarily indicate subadult age. Weconclude that, together with other compelling evidence,given the metatarsal fusion pattern of OH 8, and thedental development of the OH 7 mandible, it is highlyunlikely that the OH 8 foot and the OH 7 mandible arefrom the same individual.

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Author indexAbdala, Fernando · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 32, 35, 37, 40, 47, 51

Alberdi, Maria Teresa · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Albrecht Manegold · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Almond, John E. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 37, 50

Angielczyk, Kenneth· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 30, 49

Antoine Louchart · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Backwell, Lucinda R. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 55

Bamford, Marion · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 30, 46, 47, 48, 52, 56

Berger, Lee · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 47, 55

Bergh, Eugene · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 31

Bever, Gabe· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Bordy, Emese M. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40, 52

Botha-Brink, Jennifer · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 31, 38, 54

Canoville, Aurore · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 31

Carlson, Kristian J.· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 32, 35, 38

Chinsamy-Turan, Anusuya · · · · · · · · · · · · · · · · · · · · · · · · 31, 32, 36, 41, 51

Christian, Andreas · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40

Cisneros, Juan C. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 33

Cohen, Brigette · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 33

Collins Cook, Della · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 35

Compton, John S. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 56

D'Elía, Guillermo · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 45

d'Errico, Francesco · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 55

da Conceição, Domingas, M.· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Day, Michael· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 34, 37, 51

de Castro Silva, Mayana· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

de Cock, Michiel · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

DeSilva, Jeremy · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57

Dirks, Paul · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39

Dowdeswell, Mark R. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 38

Dumont, Maïtena · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 34

Durand, Francois· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 35

Fernandez, Vincent· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 32, 35

Fiorillo, Anthony R. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 32

Fröbisch, Jörg · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Gastaldo, Robert · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 50

Gauthier, Jacques· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Gess, Robert · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 35

Gomez, Santiago · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Götz, Annette E. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 36

Govender, Romala· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 36

Güven, Saniye · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 34, 37

Hancox, John · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46

Henry, Amanda · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 47

Houssaye, Alexandra · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 37

Hsiang, Allison· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Hübner, Tom · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 51

Huttenlocker, Adam · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 38

Iturra-Cid, Myriam · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 45

Jashashvili, Tea· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 38

Jasinoski, Sandra · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Jinnah, Zubair · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39, 50

Jirah, Sifelani · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39

Kammerer, Christian F. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Keeling, Rachelle · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39

Kibii, Job · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 33

Klein, Nicole· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40

Knoll, Fabien · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Kostka, Aleksander · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 34

Kruger, Ashley · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40

Krummeck, William · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40

Krupandan, Emil· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 41

Kupczik, Kornelius · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 54

Labandeira, Conrad · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 41

Lewis, Patrick · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Lindgren, Johan· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 37

Looy, Cindy · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 50

Lyson, Tyler · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Marchi, Damiano· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 38

Mariscano, Claudia A. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Martinez-Maza, Cayetana · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Matthews, Thalassa· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

McKay, Ian · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 44

McPhee, Blair · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 44

Mocke, Helke · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 45

Montoya, Germán· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 45

Mostovski, Mike · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 48

Nalla, Shahed· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46

Nesbitt, Sterling· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 49

Neumann, Frank H. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46, 52

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Nicoletti, Natasha · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46

Norton, Luke · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 47

Nshimirimana, Robert · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 38

Odes, Edward · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 47

Olayiwola, Moshood · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 48

Ovechkina, Maria · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 48

Parkinson, Alexander Haig · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 48

Peecook, Brandon · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 49

Penn-Clarke, Cameron · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 49, 50

Prado, Jose Luis · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43

Prevec, Rose · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 41, 50

Redelstorff, Ragna· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 51

Richter, Martha · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29

Roberts, David · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Roberts, Eric · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 39

Rubdige, Bruce · · · · · · · · · · · · · 37, 29, 32, 33, 34, 35, 39, 40, 46, 47, 50, 51

Ruckwied, Katrin· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 36

Sanchez, Sophie· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 51

Sander, P. Martin · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 40, 53

Scarborough, Matthew E. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Scheyer, Torsten· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 42

Sciscio, Lara · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Scott, Louis · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Shelton, Christen D. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 53

Sidor, Christian · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 30, 38, 49, 55

Smith, Roger M.H. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 29, 56

Smith, Roger· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 30, 31, 49, 54, 55

Steininger, Christine · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 54

Steyer, Jean-Sebastien· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 30, 55

Stynder, Deano D. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 43, 54

Tafforeau, Paul · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 32, 35

Thackeray, Francis · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 55

Thomas, Daniel B. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 32

Tocheri, Matthew · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57

Tsikos, Hari · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 52

Tsuji, Linda· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 55

Tumarkin-Deratzian, Allison R. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 32

Tütken, Thomas· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 34

Val, Aurore · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 33, 55

Van Arsdale, Adam · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57

Van Dijk, D. Eduard · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 56

Viglietti, Pia A. · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 56

Weiss, Elizabeth · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57

Welman, Johann · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 57

Yates, Adam · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 35, 46, 49

Zipfel, Bernhard · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 46, 57

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Preface to Professor Nina Jablonski’s public lecture on“Skin: Its Biology in Black and White”

The Palaeontological Scientific Trust (PAST) has regularly presented a keynote scientific lecturefor the public, in fulfilment of its mission to preserve, promote and protect Africa’s ancientheritage, and as part of its mandate to integrate education, research and outreach in the originsciences (palaeontology, palaeo-anthropology and archaeology). The public lecture, which fallsunder PAST’s Public Understanding and Engagement Programme, has since its inception beensupported by the Standard Bank, who through PAST remains a founding and major supporterof the origin sciences. The lecture has gained tremendous popularity due to its high-profilescientific speakers presenting complex scientific information in an understandable manner.Previous speakers include Berhane Asfaw, Ron Clarke, Richard Dawkins, Richard Leakey,Paul Sereno, Phillip Tobias and Tim White.

In honour of the late Phillip Tobias, PAST’s founding Scientific Advisor and more recentlyits Scientific Patron, the 2012 lecture is the first in the series to be renamed the Phillip TobiasMemorial Lecture. As a reflection of PAST’s efforts to promote the social relevance of the originsciences, we had endeavoured to make this lecture feature a scientific topic that comments onthe complex and challenging social issues facing humanity today. It is therefore fitting that thelecture commemorates not only Phillip Tobias’ scientific prowess, but also his life-long advocacyof human rights and social justice.

The 8th Standard Bank/PAST Keynote Lecture was presented by scientist and author, Profes-sor Nina Jablonski, at the Soweto Theatre, Johannesburg, on 19 September 2012. Prof. Jablonski,of the Anthropology Department, Pennsylvania State University, U.S.A., is a biologicalanthropologist and head of her department. She is well-known for her research on the evolu-tion of skin colour, and she has published numerous papers and books on primate and humanevolution. She conducts fieldwork in Asia and Africa, and in 2002 was elected a Fellow of theAmerican Association for the Advancement of Science. She is also an associate of theStellenbosch Institute for Advanced Study. Her PAST keynote lecture was entitled ‘Skin: ItsBiology in Black and White’ and was delivered to an audience in excess of 350 individuals. RickMenell, Chairman of PAST, served as the Master of Ceremonies, and Derek Hanekom, the thenDeputy Minister of Science and Technology (promoted to the post of Minister in this Depart-ment in October 2012), delivered opening remarks.

PAST would like to thank Prof. Jablonski, and her collaborator and husband, Dr GeorgeChaplin, for their efforts to help the organization promote the understanding and relevance oforigin sciences in the modern world. Prof. Jablonski’s lecture underscored the message of ourcommon human origins in Africa, a message that PAST sees as core to its mission and the sevenprogrammes it supports. It is individuals and institutions such as Prof. Jablonski and theStandard Bank, that ensure PAST’s ability to teach the youth of Africa and indeed the world,that they can proudly proclaim Africa to be home.

To celebrate the broader pan-African mission of PAST since the 2011 launch of its Scatterlingsof Africa project, PAST wanted to ensure the more widespread distribution of the lecturecontent through its publication in an African-based academic journal. Given PAST’s longstand-ing support of Palaeontologia africana, this journal was the preferred choice for the first publica-tion of the Standard Bank/PAST keynote lecture. Copies of this article will be disseminatedthrough PAST’s foundational education programme, the Walking Tall Schools Project, to educa-tors, learners and members of the public lacking access to the journal. For information on PASTand the seven programmes it supports, please refer to www.past.org.za.

Andrea LeenenCEO, PAST, P.O. Box 203, Parklands, Johannesburg, 2121 South Africa

&Robert J. Blumenschine

Chief Scientific and Educational Strategist, PAST, and Professor of Anthropology,Rutgers University, New Brunswick, New Jersey, U.S.A.

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Standard Bank/PAST First Phillip Tobias Memorial Lecture

Skin: Its Biology in Black and WhiteNina G. Jablonski

Department of Anthropology, 409 Carpenter Building, The Pennsylvania State University, University Park, PA 16802, U.S.A., andStellenbosch Institute of Advanced Study (STIAS), 19 Jonkershoek Road, Stellenbosch, 7600 South Africa

E-mail: [email protected]

Human skin comes in a wide range of sepia tones, from darkestbrown to near ivory, but not in black or white. The sepia rainbowof skin colours grades from dark to light, from the equator to thepoles. The geographic distribution of skin pigmentation hasbeen known and studied by many people over nearly three cen-turies, but only in the last 20 years have new kinds of data be-come available that allow skin colour variation to be wellunderstood. Skin pigmentation is a topic where ‘the rubbermeets the road’ because it is at the meeting point of evolutionarybiology and human experience. Scientists and the general publicalike are engaged by skin colour diversity, and the social sensitiv-ity that once inhibited both scientific research and social dis-course on the topic has now ebbed. People want to know howskin colour evolved and why it has come to have the meaning ithas in modern societies.

One of the biggest challenges in studying the evolutionof skin is that there is very little actual preserved ancienthuman skin to study. There is no fossilized human skin yetconfirmed, only the skin associated with mummified,frozen, or bog-preserved corpses dating to no more than afew thousand years old (Jablonski 2006). The study of skinand skin colour evolution in humans thus relies on evi-dence from comparative study of the anatomical andphysiological characteristics of the skin of living peopleand primates, along with the comparative study of thegenes that determine these characteristics. A lot of infor-mation has been inferred about the nature of the skin andhair in the last common ancestor (LCA) of humans andchimpanzees, for instance, by examining these featuresand some of the genes associated with them in humans,apes, and monkeys (Jablonski 2004). This examinationshows that the LCA probably had dark hair coveringlightly pigmented skin, very much like a modern chim-panzee. Exposed skin on the face and hands would havedarkened by tanning when the skin was exposed tostrong sunlight, but the skin covered by hair would haveremained untanned. We can extend the same inference tothe appearance of the skin in one of the best known the

early fossil hominin species, Australopithecus afarensis,known from middle Pliocene sites in Ethiopia and Tanzania.The skeletal evidence of A. afarensis indicates that, althoughthe species was habitually bipedal, it was not an energeticbipedal strider and runner. Rather, the species had ape-like body proportions, retained abilities to climb trees, andprobably was not a long-distance runner or fast walker.These aspects of hominin lifestyles are important becausebody temperature is elevated during vigorous exercise,and problems of dissipation of heat can become criticalespecially when hominins are exercising in hot environ-ments. The likely cause of the loss of most functional bodyhair in hominins was the need for more efficient coolingfrom evaporation of sweat from the surface of the body(Jablonski 2004). Evaporation of expressed sweat worksextremely well if the skin is naked, but not well if the skinis covered with dense hair (Folk & Semken 1991). Whenthe skeletons of early members of the genus Homo arecompared with those of Australopithecus, the differencesbetween the two in the likely modes of locomotion areimmediately apparent (Crompton et al. 1998; Ruff 2008;Ruff 1991). Unlike Australopithecus, early Homo was abipedal strider and runner (Bramble & Lieberman 2004).These activities raised metabolic levels and generatedbody heat that had to be liberated from the body’s surface.Dissipation of excess body heat probably was accom-plished primarily by evaporation of sweat, and thesweat was produced by a high density of eccrine sweatglands on the body’s surface (Jablonski 2004). Keepingcool is of particular importance in highly encephalizedprimates: the brain is sensitive to elevated core tempera-ture. Eccrine sweat evaporating on nearly naked skincreates a whole-body cooling system that helps to main-tain the brain’s thermal homeostasis during intense exer-cise (Caputa & Cabanac 1988; Shibasaki & Crandall 2010),and enhanced body cooling through sweating nodoubt released a physiological constraint that made

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Human skin comes in a range colours, grading from very dark brown near the equator to near ivory near the poles. This colour gradientresulted from the action of natural selection. The original skin tone for members of the Homo lineage, including Homo sapiens, was darkbrown. Diversification of skin colouration occurred as some Homo sapiens populations dispersed to regions of lower ultraviolet radiation(UVR), mostly outside of Africa. Humans who began to inhabit higher latitudes faced reduced opportunities to make vitamin D in theskin because of lower and more seasonal levels of UVB from ambient sunlight. Depigmented skin evolved under these conditions, andgenetic evidence indicates that mutations leading to loss of melanin pigmentation occurred multiple times in hominin history whengroups invaded regions of lower UVB. Because similar skin colours have evolved multiple times under the same environmentalconditions, classification of humans according to skin colour is fraught with problems. The colour-based races defined in the 18thcentury are congeries of physical and behavioural traits that do not exist. The persistence and propagation of these concepts has causedserious problems in many places, and can only be countered by effective education.

Keywords: evolution, natural selection, pigmentation, melanin, eumelanin, ultraviolet radiation, UVB, folate, vitamin D,depigmentation, convergent evolution, race.

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possible increased brain size in the human lineage(Jablonski 2010a).

The evolution of functional nakedness was not the onlymajor change that occurred in the early evolution of thegenus Homo. Naked skin was vulnerable to the damagingeffects of ultraviolet radiation, and so we reasoned fromfirst principles that permanent dark pigmentation in theskin would have had to have evolved in early Homo at thesame time as hairlessness in order to protect individualsfrom these effects (Jablonski & Chaplin 2000). This infer-ence was later supported by genetic evidence indicatingthat the form of the MC1R pigmentation that producespermanent dark pigmentation in darkly pigmentedAfricans today had been invariant for as long as 1.2 millionyears (Rogers et al. 2004). Early members of the genusHomo living under equatorial or near-equatorial sunlightcontaining abundant ultraviolet radiation (UVR) were,thus, hairless and darkly pigmented (Jablonski & Chaplin2000).

The pigment melanin, specifically, the intensely darkbrown form of melanin called eumelanin, is a naturalsunscreen that confers superior protection against thedamaging effects of UVR (Brenner & Hearing 2008; Kolliaset al. 1991). The intensity of UVR at the earth’s surface is re-lated to latitude and humidity, with arid areas near theequator having the highest levels (Chaplin 2004). Outsideof the tropics, UVR levels decline significantly, and theamount of UVR that falls on any particular area dependson the energy of the radiation emitted by the sun. Themost biological significant types of UVR that fall on theearth are long wavelength UVR called UVA and mediumwavelength UVR called UVB. The more energetic of theseis UVB and it is readily absorbed and scattered by oxygenand ozone in the atmosphere. Outside of the tropics,when the sunlight passes obliquely through the atmo-sphere during the winter months, little or no UVB reachesthe earth’s surface, while most UVA passes through theatmosphere with visible light (Madronich et al. 1998).Ultraviolet radiation is mostly harmful to living systemsbecause it damages DNA, cell membranes, and connectivetissues, and biological systems have evolved elaboratemechanisms to repair damage caused by UVR (Cleaver &Crowley 2002; Rothschild 1999). For vertebrate animals,one of the most effective means of preventing damagefrom UVR is eumelanin pigment in the integument.When incorporated into the epidermis of the skin,eumelanin absorbs and scatters most UVB, thereby spar-ing damage to deeper layers including the blood flowingthrough the cutaneous blood vessels (Brenner & Hearing2008; Tadokoro et al. 2003; Zmudzka et al. 2006).

It was long thought that dark, eumelanin-rich pigmen-tation originally evolved in humans in order to protectagainst skin cancer caused by damage to DNA in the skin.When it was demonstrated that this was unlikely becausemost skin cancers afflict people after their reproductiveyears (Blum 1961), a rethinking of the role of eumelaninwas required. A key discovery was published in 1978,when it was shown that the B vitamin folate was sensitiveto ultraviolet radiation (Branda & Eaton 1978). Folate is avitamin that is obtained from green leafy vegetables,

citrus fruits, and whole grains, and is of critical importanceto health because it is necessary for making and repairingDNA (Lucock & Daskalakis 2000). Folate sufficiency isimportant for normal reproduction because it is a precon-dition for healthy sperm production in men and fornormal development of early embryos in women (Bower& Stanley 1992; Fleming & Copp 1998; Mathur et al. 1977).Folate deficiency is one of the main causes of the family ofbirth defects called neural tube defects (Bower & Stanley1992; Fleming & Copp 1998). Because of the important roleplayed by folate and related compounds in successfulreproduction, we reasoned that protection of these impor-tant biomolecules was the primary cause for the evolutionof permanent eumelanin protection in the skin (Jablonski& Chaplin 2000). Eumelanin-rich skin protected allmembers of the genus Homo, from the species of the EarlyPleistocene through Homo sapiens. The evolution of theHomo lineage leading to anatomically modern humans,Homo sapiens, occurred in Africa, and all members of thislineage were darkly pigmented (Jablonski & Chaplin 2000).The question then arises as to how and when the modernpattern of skin colour diversity arose. For the answer tothis question, we must examine the pattern of dispersalsof modern people outside of tropical Africa, and the UVRconditions that obtained in these environments.

Modern humans, Homo sapiens, evolved in Africa between200 000 and 120 000 years ago, and the first half of ourspecies existence was spent only in Africa. During thistime, populations expanded and dispersals into theextremities of the continent proceeded, accompanied byextensive genetic diversification (Tishkoff et al. 2009). Thechanges that occurred in human populations during thisperiod would have included subtle changes in pigmenta-tion, as populations adapted to local UVR conditions orexperienced genetic bottlenecks due to small populationsizes. The genetic basis for evolutionary changes in pigmen-tation among modern people living in Africa is not wellunderstood. Approximately 80 000 years ago, small popu-lations of modern people left Africa, taking with them asmall fraction of the continent’s genetic diversity. Thesepeople were culturally sophisticated and their move-ments – into southeast and east Asia first, then later intoEurope – occurred quickly by sea and over land (Stringer2000). These movements saw people entering highlydiverse solar regimes, including those with very lowlevels of UVR. A large fraction of the land mass of Eurasialies at high latitudes which receive low UVR, and particu-larly low UVB (Chaplin & Jablonski 1998). As peopleentered these lands, they were subjected to lower andmore strongly seasonal levels of UVR than they had everexperienced, and this had profound effects on humanhealth (Chaplin & Jablonski 2009).

Ultraviolet radiation is not a universally malign influ-ence. Its single most important positive action is initiationof the process of vitamin D formation in the skin by UVB.This process occurs in all terrestrial vertebrates and isessential for life (Holick 2003). Vitamin D is necessarybecause it permits absorption of calcium from the diet.Vitamin D deficiency is a serious problem that leads toskeletal malformation and weakness that manifests itself

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as rickets in children and osteomalacia in adults (Dimitri &Bishop 2007). Serious and persistent rickets in youngwomen leads to pelvic deformities that impair or preventnatural childbirth, so vitamin D deficiency can directlyimpact reproductive success. Because vitamin D is alsoresponsible for maintenance of the health of the immunesystem, vitamin D deficiencies are related to increasedsusceptibility to infectious diseases, certain cancers, andautoimmune disorders (Holick 2005; Ponsonby et al. 2005)(Holick 2007; Holick & Chen 2008).

On the continent of Africa, the region experiencing thelowest and most seasonal levels of UVB is the southerncoast of South Africa. At and near the winter solstice, thereis no UVB in the sunlight over this region, and no vitaminD production can take place in human skin, regardless ofskin colour (Jablonski & Chaplin, unpubl. obs.). Theabsence or attenuation of UVB reaching middle and highlatitudes during the winter months had profoundinfluences for the evolution of skin pigmentation. Becauseeumelanin is a superior sunscreening agent, and becauseUVB is necessary to make vitamin D in the skin, it wouldhave been beneficial for people living under conditions ofweak and highly seasonal sunlight to have less eumelaninin their skin (Jablonski 2004; Jablonski & Chaplin 2000;Loomis 1967; Murray 1934). In any given period of time,more vitamin D precursor can be in lightly pigmentedskin than in darkly pigmented skin, and so lighter skinwas favoured by natural selection under low UVB condi-tions (Norton et al. 2007; Webb & Holick 1988). Light skin isactually depigmented skin, and genetic mutations lead-ing to depigmentation evolved independently in theancestors of modern east Asians and modern westernEuropeans (Lamason et al. 2005; Norton et al. 2007). Theimportance of vitamin D production in the skin is attestedby the fact that genetic fixation of the pigmentation genemutations leading to loss of pigmentation occurredthrough selective sweeps resulting from positive selection(Norton et al. 2007). Loss of eumelanin pigmentation hasalso been inferred to have occurred earlier and independ-ently in Homo neanderthanlensis, the distant cousin of Homosapiens that inhabited low and moderate UVB environ-ments Europe and the circum-Mediterranean (Lalueza-Fox et al. 2007).

The convergent evolution of light pigmentation on atleast three occasions in hominin history – in the ancestorsof modern western Europeans, modern east Asians, andNeanderthals is remarkable, and has important conse-quences for the use of skin colour as a feature in humanclassification. Similarly, the evolution of repigmentation isequally important. As people have moved around in thelast 20 000 years, some depigmented populations havere-entered regions with strong UVR and have undergonegenetic mutations leading to enhanced tanning abilitiesand darker baseline pigmentation. This has been demon-strated for some New World populations (Quillen 2010)and is strongly suspected for some groups that nowinhabit southern reaches of the Indian subcontinent.

Through time, people have become increasingly mobileas the result of innovations in transportation technology,spurred on by interests in distant resources and trade.

This has resulted in people moving around over longerdistances at faster speeds, especially in the last 500 years.The mismatches between skin pigmentation and environ-mental UVR that have arisen because of these rapid move-ments have resulted in profound and mostly negativeeffects on health (Chaplin & Jablonski 2009; Jablonski &Chaplin 2012). People living in earlier times were less mobile,and would not have been exposed to solar regimes signifi-cantly different from those experienced by their ances-tors. But this situation is common for modern people,whether they are vacationing or are living permanently ina region distant from their ancestral homeland. Even forpeople who have not moved far in recent millennia, thereis often a mismatch between skin pigmentation and sunexposure because of indoor lifestyles in cities and a lack ofsun exposure. Lightly pigmented people living in highUVR environments face significantly elevated risk of skincancer (De Gruijl 1999; Rees 2000; Rigel 2008). Darkly pig-mented people living under low or seasonal UVB condi-tions face an elevated risk of vitamin D deficiency(Brunvand & Haug 1993; Fogelman et al. 1995; Fonsecaet al. 1984). Vitamin D deficiencies are also increasinglycommon among city dwellers of all skin colours whospend most of their time indoors, or among people whoroutinely wear concealing clothing and chemical sunscreenwhen outdoors (Chatfield et al. 2007; Davies et al. 1986;Forrest & Stuhldreher 2011; Le Goaziou et al. 2011).Although relatively few epidemiological studies on vita-min D status have been conducted in South Africa, prelim-inary indications are that vitamin D deficiency is commonand has serious consequences for disease susceptibility(Martineau et al. 2011).

Skin is the interface between the body and the physicalenvironment, and skin pigmentation evolved to regulatethe penetration of UVR into body under different envi-ronmental conditions. The gradient of skin pigmentationseen in modern people, from darker tones near the equatorto lighter ones closer to the poles, is the product of twocompeting clines, one favouring photoprotection in areasof high UVR, the other favouring photosynthesis ofvitamin D in areas of low UVR (Chaplin & Jablonski 2009;Jablonski 2010b; Jablonski & Chaplin 2010). People withmoderate pigmentation living in middle latitudes (betweenabout 23� and 37�) generally have good abilities to tan,thereby gaining a measure of increased protection fromsummer sun. Skin pigmentation is a superb example ofnatural selection acting on the human body, and shouldbe used as a model system for teaching evolution at alllevels of the curriculum.

What skin colour does not provide is a unique marker ofancestry for purposes of classification of people intogroups. Visibly identical skin phenotypes have evolvedfrom distinct genotypes that arose from independentmutations. This means that skin colour itself cannot beused as a unique indicator of genetic ancestry or as acharacteristic for sorting people into races because thesame colour is associated with multiple ancestries andraces. We recognize this now, but this fact was not knownto the naturalists who composed the first classifications ofhumans and the philosophers who first defined races.

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When Carolus Linnaeus created the first scientific classifi-cation of humans, he placed them into four groups, sortedby skin colour and continent. The variations of Homo henamed in 1748 in the 6th edition of his Systema Naturaewere Europaeus albus (white), Americanus rubescens (red),Asiaticus fuscus (brown), and Africanus niger (black).Linnaeus had limited information on which to base aclassification of humankind, and so this simple classifica-tion must not be judged too harshly. What is significant iswhat happens in subsequent editions of the SystemaNaturae. By the 10th edition of 1758, Linnaeus hadchanged his classification to include more characteristicsof each group. Physical traits other than skin colour, suchas eye colour and hair texture were added, but he alsoincluded behavioural attributes. Europeans weresanguineus (sanguine), [Native] Americans were cholericus(choleric), Asians were melancholicus (melanchology), andAfricans were phlegmaticus (phlegmatic). He also charac-terized the four varieties of people according to regitur orhow they were governed, so that Europeans weredescribed as being ruled by ritibus (rites), Americans byconsuetudine (custom), Asians by opinionibus (opinion),and Africans by arbitrio (will). Linnaeus was a naturalistand Systema Naturae was considered the first ‘scientific’classification of living things. It was extraordinary, there-fore, that Linnaeus departed from his usual custom of de-scribing organisms on the basis of only their physicalattributes to describe people on the basis of dispositionsand cultural behaviour as well as physical traits (Broberg1983). The mixing of physical and behavioural traits thatLinnaeus introduced heralded a new era in human classi-fication, and presaged the first description of races.

Linnaeus’s classifications of humans were widely readand discussed by scholars of his day. Other naturalists andphilosophers took up the challenge of human diversity,and by the late 18th century, no less a figure than ImmanuelKant had published his own classification of humans.Kant was the first person to place people into races (Rassenin German), which were defined by skin colour and placeof origin (Bernasconi 2001; Shell 2006). Kant placed greatimportance on place of origin because climate (and sunexposure) had purposeful influences on physical traits –notably skin colour – and on the collective motivationexhibited by a race and the level of civilization it couldattain (Bernasconi 2006). Kant considered races and racialcharacteristics to be fixed and immutable, and on thispoint he clashed with many of his contemporaries(Bernasconi 2006). He was also the first to rank racesclearly and unambiguously: ‘Humanity is at its greatestperfection in the race of the whites. The yellow Indians dohave a meagre talent. The Negroes are far below them andat the lowest point are a part of the American peoples’(Bernasconi 2002).

The most significant aspect of Linnaeus’s and Kant’staxonomies in the context of this paper is that bothLinnaeus and Kant considered that the colours of skinthey recognized were unique to geographical groups ofhumans, and that skin colour was unerringly connectedwith other physical and behavioural traits. They conceivedof defined varieties or races of humans as discrete pack-

ages of biosocial attributes. These conceptions have lastedand have been propagated, despite the accumulation oftwo centuries of evidence showing that discrete races donot exist. The study of skin pigmentation evolution hasrevealed that similar skin colours evolved independentlymany times in hominin history, and that pigmentation isnot, for the most part, genetically connected to otherphysical traits. Further, there is no genetic relationshipbetween physical, behavioural, and cultural traits. Thus,the original conception of colour-based human raceswhich many societies have operated under for over 200years is flawed and baseless.

Skin colour is a trait of great biological significance thatillustrates action of evolution by natural selection on thehuman body. It is also a trait that has been used, incor-rectly and persistently, for classifying people into groupswhich were considered to be elevated or diminished instatus because of colour. This unfortunate state of affairscan be changed, if there is motivation and a commitmentto change through education. We are one people, and skincolour unites us, not divides us, through the process ofevolution.

This paper is based upon a transcription by Ann Smilkstein of the 8th PAST/Standard Bank Annual Lecture I delivered on 19 September 2012 at the SowetoTheatre. I am very grateful to PAST and to Standard Bank for inviting me to givethis lecture. With the renaming of the lecture series, I am especially honoured tohave been invited to give the 1st Philip V. Tobias Memorial PAST/Standard BankAnnual Lecture. Andrea Leenen, Robert Blumenschine, and Ann Smilkstein ofPAST masterfully organized the lecture and the events surrounding it. My stay inSouth Africa was made possible by a Guggenheim Fellowship and by a fellowshipfrom the Stellenbosch Institute for Advanced Study. My deep thanks go to mysenior research technician, Tess Wilson, for continuing support with research andpresentation materials, and to George Chaplin for many fruitful discussions,ongoing collaboration, and comprehensive support.

REFERENCESBERNASCONI, R. 2001. Who invented the concept of race? Kant’s role in

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CROMPTON, A.W. 1962. On the dentition and tooth replacement in twobauriamorph reptiles. Annals of the South African Museum 46(9), 231–255.

HOPSON, J.A. & BARGHUSEN, H.R. 1986. An analysis of therapsid relationships.In: Hotton, N., MacLean, P.D., Roth, J.J. & Roth, E.C. (eds), The Ecology and Biologyof Mammal-like Reptiles, 159–168. Washington, Smithsonian Institution Press.

ROMER, A.S. 1966. Vertebrate Palaeontology (3rd edn). Chicago, University ofChicago Press.

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VAN DER MERWE, N.J., LEE-THORP, J.A., THACKERAY, J.F., HALL-MARTIN, A.,KRUGER, F.J., COETZEE, H., BELL, R.H.V. & LINDEQUE, M. 1990. Source-areadetermination of elephant ivory by isotopic analysis. Nature 346, 744–746.

PERIERA, L.M. 2009. The Wits Online Phytolith Database. University of theWitwatersrand, Johannesburg, South Africa. Online at: http://web.wits.ac.za/Academic/Science/GeoSciences/BPI/Research/WOPD/ (accessed 15 February2010).

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